Reclassifying Reptiles

When I was a child, my favorite animals were reptiles and amphibians.  I spent countless hours collecting toads under the street lights at night, chasing whiptails in the sun, racing to grab spiny lizards before they darted up the tree, all while seeking the holy grail of childhood herping: snakes.  

Reptiles and amphibians just seemed to fit together.  Of all the terrestrial chordates, they alone were cold blooded.  They were discussed in tandem books and documentaries, and they were even lumped together under the scientific discipline of herpetology. 

Herpetology is a discipline that studies an arbitrary assemblage of animals. Illustration by author.

Something about these magical creatures captivated my young mind. Years before I ever met my first scorpion or tarantula, I had known my first love. 

Reptiles and amphibians tend to elicit extreme reactions in people.  People seem to either hate, say frogs or snakes, or love them with little middle ground.  Collectively reptiles represent some of the strangest, and yet most familiar of all animal groups.  Love them or hate them, everyone knows a snake when they see it.  

However reptiles are not limited to the modern familiar forms, they also include the famous dinosaurs that every young child spends a period obsessing over.  

Reptiles, as we normally think of them, are not so easily classified as “cold blooded scaly things that aren’t fish.”  To see why, we should start by looking at how reptiles were initially classed in the Linnaean system, and how our understanding has grown over time.  So join me, as we explore life in endless forms most beautiful.

In the last entry in this series on taxonomy, I introduced the concept of monophyletic clades for classification, in contrast to the historically traditional Linnaean ranks. Today, I want to look at the history of the classification of reptiles and amphibians, and begin to take a closer look at the reptile family tree.

When Linnaeus published the first edition of Systema Naturae in 1735, he had initially lumped reptiles and amphibians together into sort of a “catch all” class he called Amphibia.  Because he lived in Sweden, which has a very low diversity of reptiles compared to the southern US where I live, he knew comparatively little about them.  

This first edition included only a handful of described reptile and amphibian species totaling 27.  Even though the Linnaean group “Lacerta” included lizards, crocodilians and salamanders, the fact that he created two additional groups for the other tetrapod herps, Rana for frogs and Testudo for turtles did reflect the growing awareness of greater diversity within the Amphibia which would later be built upon.  

The Kingdom Animalia of the fist edition of SI. Linnaeus had set out to document each species, believing this to be a straightforward and finite task, after all Adam had done this hadn’t he? He could not have known, nor could he have imagined what he was getting into.

As a side note, it turns out that Linnaeus intensely disliked the animals he grouped as the amphibia, often ironically expressing gratitude that they had a low diversity compared to what he saw as more noble classes like mammals and birds.  He stated in the 1735 introduction to the class: “The Creator in his benignity has not wanted to continue any further the Class of Amphibians; for, if it should enjoy itself in as many Genera as the other Classes of Animals, or if those things were true that the Tetralogists have fabricated about Dragons, Basilisks, and such monsters, the human genus would hardly be able to inhabit the earth.” 

Also note from that quote his distaste for mythological animals.  This is an important part of Linnaeus’ mindset that was deeply influential on the field of biology, more on that in another entry.

By 1740, with the second edition published Linnaeus had divided the Amphibia into two orders; the Reptilia or the amphibians with legs; which included frogs, turtles, salamanders, lizards, and crocodilians; and the order of legless amphibians he called Serpentes which included snakes, as well as legless amphibians.

Eight years later, the sixth edition (which was technically the third edition directly authored by Linnaeus) saw a dramatic increase in diversity among the described amphibians, including 6 new genera, and 40 new species.  

Notably, by this time, Linnaeus had developed something of a love-hate relationship with snakes that would last the rest of his life. Having both worked as a physician and dedicated some time to attempting to find new remedies for snake bites, Linnaeus apparently developed a fascination with snakes. By the sixth edition, he now drew a clear distinction between Colubrids and Vipers.

Finally, in 1758, after spending a few years traveling, lecturing and further refining his work, Linnaeus would publish the largest update to Systema Naturae to date, and the single greatest increase for the class amphibia in the entirety of his work.  The 10th edition of Systema Naturae would add another 117 described amphibia to the list, as well as introducing another order for a handful of cartilaginous fish he called Nantes.

In it Linnaeus reveals that his opinion has not changed much regarding the class amphibia, stating: “These most terrible and vile animals are distinguished by their unilocular and single chambered heart, arbitrary lungs, and divided penis. Most amphibians are rough, with a cold body, a ghastly color, cartilaginous skeleton, foul skin, fierce face, a meditative gaze, a foul odor, a harsh call, a squalid habitat, and terrible venom. Their Author has not, therefore, done much boasting on their account.”  I rather disagree, but to each their own I suppose.

The final edition of Systema Naturae authored by Linnaeus was a large update to the work as a whole, but relatively unremarkable in terms of his treatment of the amphibia.

Linnaeus had made some interesting insights in his work of classifying the natural world. In the first edition, whales were grouped with fish, but he corrected this in the 10th.  He was also one of the first to remove bats from being classed among the birds because he recognized their mammalian nature.  

But for all his insight, he seemed only dimly aware of the fundamental natures of what we now call reptiles and amphibians.  To be fair, Linnaeus was primarily a botanist as his main area of expertise.  

Still, the Linnaean system had treated the amphibia largely arbitrarily, basing the classifications on the number of limbs more than anything else.  For example, in 1776 naturalists were perplexed by the existence of an animal that defied categorization in his normal treatment of the animal world, the Siren.

Such was the fascination with the animal that some of Linneaus’ correspondences were printed and distributed as a dissertation on the siren.  Linnaeus describes it thus: “It is shown to us how the animal diverts from the general characterizations in the Animal kingdom; in lack of hair and feathers it can be neither Mammal nor Bird; among Amphibia it cannot be assigned to any of the known divisions, neither to the Reptilia, which have four feet and no gills; nor to the Snakes, that are without feet, fins and gills; nor to Nantes, which all have fins. In view of this, Author shows the necessity of establishing a new Order in Reptilia, with two feet, lungs and gills. Many would think this animal is simply a larva of Lacerta Iguana or another, but both claws and its sound deny this…”

His reference to the larva of an unambiguous lizard reveals that Linnaeus had not truly understood the difference between salamanders and lizards, like iguanas.  

Salamander and lizard. While both share the same “lizard shape” at first glance, they are fundamentally different creatures. The squamate on the right has more in common with bears and parrots than it does with the salamander. This “lizard shaped body” is actually the basic body shape of all tetrapods (including humans) and their association with each other is merely superficial. Images from Wikimedia commons, edited by author.

He was aware that some of the amphibia undergo metamorphosis, and others are born or hatched as miniature adults, but it would seem he had not developed an appreciation for the implications here.

The distinction in reproductive modes of the group was noted in the introduction to the tenth edition as follows: “A polymorphous nature has bestowed a double life on most of these amphibians: granting that some undergo metamorphosis and others cast off their old age. Some are born from eggs, whereas others bear naked young. Some live variously in dry or wet, whereas others hibernate half the year. Some overcome their prey with effort and cunning, whereas others lure the same prey to their jaws as if by magic.”  

Despite this awareness of the dramatic difference in reproductive processes, the distinction between the Rana and the Lacerta was the presence or absence of a tail.  Ranids, or frogs, were naked (scaleless), four legged reptiles that also lacked tails, and Lacerta, defined by having four feet and tails, were collectively called lizards.  This included both the crocodilians and salamanders.

The latter were considered to be a subset of “Naked aquatic lizards with unarmed or un-clawed feet.”  For all his detailed and intricate descriptions, Linnaeus’ treatment of the class amphibia remained superficial. 

For many naturalists, the amphibia were just a collection of ugly and unappealing animals of varying levels of venom, disease, superstition, and were generally seen as varmints.  

The biggest reason for this was the human tendency to see the world anthropocentrically.  Of all the terrestrial animals, the amphibia are the least like us.  Cold blooded, not soft and cuddly like birds and mammals, they seem the most alien to us despite being the majority of terrestrial chordate diversity.     

But it is the amphibia that have more to teach us about ourselves than any other animal group.  In learning about them, we came to understand what we are, and to see our true place in the world.  

So what does distinguish what we now call reptiles from amphibians?  To answer this question, let’s continue tracing the history of their classification.

The name reptile derives from the Latin word for crawling or creeping.  In Linnaeus’ time the name was interchangeable with amphibian.  While Linnaeus used the name amphibian, it is important to keep in mind that Reptile was the more popular term among French naturalists, but both referred to the class Amphibia during this period.  

At the beginning of the 19th century a French geologist by the name Alexandre Brongniart published a paper titled Testing for a natural classification of reptiles in which he recognized the dramatic differences that set the frogs and salamanders apart from the rest of the reptiles, and proposed a new name for them: Batrachia.  

Left: Alexandre Brongniart. Right: Pierre Andre Latreille. Wikimedia commons and public domain.

(As a side note, frogs and salamanders actually do form a monophyletic clade within the amphibians and that clade still bears the name Batrachia to this day.)  

A couple of years later a prominent entomologist by the name of Pierre Andre Latreille, would concur with this classification, and would opt to classify the Batrachia apart from reptiles entirely, making him one of the first to fully comprehend that these are fundamentally different groups.  

As things played out, the new class would take on the name amphibia, as distinguished from the Reptilia which contains the remainder of Linnaeus’ original class, minus the cartilaginous fishes sometimes included.  This brings us to the modern arrangement of reptiles and amphibians as wholly separate categories we are used to.

Of course, such a large taxonomic revision would take time to become widely used, but the amphibian class had some powerful advocates including Thomas Huxley, and Earnest heckle.

Left: Thomas Huxley Right: Ernst Haeckel. Wikimedia commons.

The distinction that Brogniart and Latreille based their revisions on, and upon which the advocates who concurred converged, was their fundamentally different mode of reproduction.  

Amphibians, as we now define them, go through an aquatic larval stage of development after hatching.  This mode of reproduction is similar to how many fish (to which the basal chordates belonged) reproduce. 

Reptiles on the other hand, are able to bypass the larval foraging stage.  Instead, the embryo is able to develop inside the egg before hatching, thus reducing the threat of predation and removing the burden from the larva of needing to seek out its own food. 

This is made possible by the presence of a membrane called the amnion.  The development of this trait made it possible to reproduce on land, far from the bodies of water that amphibians depend on.  

General anatomy of the amniote egg. Illustration by author.

More importantly, it is a trait shared with all other terrestrial chordates, minus the amphibians.  This meant that reptiles were more closely related to mammals and birds than to amphibians.

This more insightful distinction in classification would find widespread acceptance among naturalists during the early 19th century.  This gives us our familiar land animal groups; mammals, birds, reptiles and amphibians.

Things became more complicated with the emergence of paleontology around the beginning of the 19th century, and even more so by a series of developments in biology, culminating in the confirmation of common ancestry for the origin of species in the latter half of the century.

With the discovery of the first dinosaurs, and the budding field of embryology, the question was being asked what the relationships were between large taxonomic groups.  

The realization that the world hadn’t always existed in its present form had come some time earlier from the field of geology, but even in Linnaeus’ time it was beginning to be realized that the same was true of the network of life as well.  

Linnaeus himself became suspicious of the assumption of the fixity of species, or more correctly the fixed number of species, as he experimented with plant hybrids near the end of his life.  He famously left out a line from his observations in the introduction to the 12th edition which assumed a constant number of extant species.  

In another work, he also wrote about a species of thalictrum that he hypothesized to be a daughter species derived from a parent and as he put it “it seems to me to be the product of its environment/the daughter of time.” 

While the conversation about evolution would become a hot topic among naturalists leading up to the point when Charles Darwin and Alfred Russel Wallace, building upon the work and insights drawn by others like them, would finally solve the puzzle of the primary mechanism of evolution, it was already known in the 19th century that some species had existed in the past, that do not presently exist. 

Left: Alfred Russel Wallace Right: Charles Darwin. Wikimedia commons.

This raised the question of how these extinct animals should be classified.  Which would have been a straightforward task if not for one thing: many of these paleo creatures belonged to groups that do not exist now.

Try to picture this.  Naturalists of the time were well familiar with the modern classes of tetrapods; birds, mammals, reptiles and amphibians.  But as more paleofauna were discovered, the question “which group is it?” had to be answered with “none of the above.”  The implications were staggering and stranger than previous conceptions of the world’s history.

A similar thing had happened to Linnaeus with the Siren, but it seems he was content to simply create a new category for the animal, without stopping to wonder why or how such a thing existed in the first place.  

However this approach of creating a new category to accommodate each new type of creature would never work for the early paleontologists.  

This was far from a handful of animals like the siren or platypus.  These were entire ecosystems of unknown groups.  The number of new classes needed would render the ranked Linnaean system far too unwieldy and awkward for any practical use.  This is the reason for the use of monophyletic clades in lieu of ranks today.  

illustration of the vastness of biodiversity which Linnaeus’ ranked system could never have accommodated, and which he could never have conceived. Illustration by author.

As the extinct species were examined and cataloged, it quickly became apparent that many of them had reptilian characteristics, but they were not anatomically modern reptiles.  Stranger still, some of these “vaguely reptilian” creatures appeared in the lineage leading to mammals.

This meant that vaguely reptilian traits were basal to the amniotes.  Even more intriguing was the discovery that the most famous of extinct animals, the dinosaurs, were not entirely cold blooded.  What’s more, some of the more derived forms had…feathers.  This meant that the ancestors of birds were also reptilian. 

Top: Dimetrodon, a “reptilish” synapsid more closely related to mammals than reptiles. Bottom: Archaeopteryx, perhaps the most famous genus of feathered dinosaurs.

At this point, it was becoming apparent that “reptile” was a poorly defined category.  It served as a “leftovers” pile for the amniotes after the mammals and birds were named.  “Reptile” just meant “an amniote that isn’t a bird or mammal.” 

Reptiles were defined not by what they are, so much as by what they are not.  They were the default for amniotes that did not belong to any better defined group.  

The realization that reptiles are actually several distinct lineages, but more closely related to each other than to mammals seems obvious in retrospect.

Snakes and lizards have so much in common and are so closely related that they pair together quite naturally, crocodilians share a number of characteristics with them and while they certainly are not lizards, they are obviously related.  

But one group of reptiles as historically defined was always different.  The testudines have a hardened carapace that forms a shell which encases most of their body.  This is a trait entirely unique to them among the amniotes.  Additionally, much of their skin lacks the scales typical of classical reptiles.  

The reptiles were initially expanded to include the newly discovered dinosaurs and other non-dinosaur paleofauna such as pterosaurs and ichthyosaurs. The definition of reptile saw several revisions during the following decades, leading into the 20th century.  

One thing was clear by the mid 19th century: mammals, reptiles and birds were all united by the amniote egg, thus revealing that amphibians were the outlier, representing a more basal state of terrestrial animals, with reptilian traits representing the next stage of development.  

Major chordate lineages that share the amniote egg as a synapomorphy. Cladogram shows a greater affinity between these lineages than any have to modern amphibians. Illustration by author.

A clear distinction was found between the ancestors of mammals, and “reptilish things.”  A synapomorphic trait that distinguishes the line leading to mammals from the rest of the known fossils at the time was the presence of a single temporal fenestra, which is an opening in the skull behind the ocular socket, the vestige of this are the temples in your skull.

Diagram of synapsid skull. Adapted from image Wikimedia commons.

By the way, a synapomorphy is a trait shared by descendant groups because it was present in a common ancestor, and thus defines the entire group (even if they lose the trait secondarily, like snakes did with legs, they are still tetrapods because their ancestors were.)

In 1863 paleontologist Thomas Huxley coined the terms Theropsids, or the “beast faced” animals and  Sauropsids, the “lizard faced” animals to describe each side of the tree.  

Although some of the early specimens on the mammal side of the tree were included with the Sauropsids in these early days, we were finally on the right track.

Around the turn of the 20th century, as more fossils were found our understanding of vertebrate evolution improved dramatically.  By this time the resolution of the Therapsid-Sauropsid split was understood in much greater detail, and the name synapsid was now widely used to describe the mammal side of the fork.  

All extant Sauropsids are a part of a subgroup called the Diapsids.  They are defined as having developed two temporal fenestra. This includes all birds, snakes, lizards, turtles and crocodilians.  

Diagram of diapsid skull. Adapted from image image on Wikimedia commons.

The diapsid tree has a fork early in its development separating the two broad groups of “reptile things” : the Lepidosaurs on one side, and the Archosaurs on the other.  

Extant diapsid clades. Illustration by author.

The Lepidosauria consist of modern squamates, rhynchocephalians, and a few extinct groups such as (possibly) ichthyosaurs and the Keuhneosauridae. 

The Archosaurs include the early branching Crocodilia, the world famous dinosaurs, pterosaurs, and of course the surviving subset of dinosaurs called birds.  

It is likely that the Archosaur branch also includes the Pantestudine clade made up of turtles and stem turtles, including plesiosaurs.  

Now that we have established a foundation for understanding the “reptile” side of the amniote tree, we can begin fleshing out the details a bit more in the next entry.  For now, let’s recap a few points. 

Remember that modern taxonomy seeks to create monophyletic groups, or a group that is defined by synapomorphies inherited from a common ancestor, and includes that ancestor and all its descendants.  

For example, the mammary organs of mammals are synapomorphic, existing only in the mammal lineage and inherited from the last common ancestor of all mammals.  

This is impossible with reptiles; because the “reptile” label describes a broader number of distinct groups than the other amniote classes.  

You can see that no matter where we draw the line when trying to define what reptiles are, we either include birds, or exclude huge numbers of what have traditionally been known as reptiles.  

Classical reptiles do not form a monophyletic group. Illustration by author.

Reptiles, as classically defined, are an invalid grouping, technically called a paraphyletic taxon because the group arbitrarily excludes some of its most recent descendants.

You may also notice that the phylogenetic position of the turtles was not shown on any of these graphics.  This is because its exact placement is still being actively researched, and this group in particular has a variety of unique historical quirks that warrant a more complete treatment later on.  

After we have fleshed out the Lepidosaur and archosaur trees, we will return to the reptile tree to bring it all together with the inclusion of the Pantestudines.  

For now, I hope you enjoyed and that I was able to entertain you with a little history and taxonomy, and that you’ll join me next time. 

Special thanks to Richard Wahlgren of the International Society for the History and Bibliography of Herpetology for compiling the majority of Linnaeus’ work related to herpetology.  

When I set out to make this entry, I was very grateful that Mr Wahlgren had already done the legwork of searching and gathering the references from among Linnaeus’ voluminous works.  

I have included a link to the paper published via ISHBH written by Mr Wahlgren below if you would like to read his overview of Linnaeus’ contributions to herpetology.  

YouTube version of this post

Kukulcania hibernalis, a childhood favorite.

One of the very first creatures to capture my attention as a child were the large, black, velvety, spiders that built messy retreats under the eaves of my back porch. Something about these animals intrigued me, and made me want to know more.

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Juvenile/subadult Kukulcania hibernalis typical of the species. Photo by C. Peterson, Lake Whales Florida, August 2021.

I didn’t know anything about evolution at the time, but I did recognize certain traits about this particular species that reminded me of tarantulas, at least from what I had seen of them in books; though it would be years before I had the opportunity to see one in person. I couldn’t articulate exactly what it was that made this spider stand out from more typical spiders, but did recognize that it had features that most other araneomorphae did not.  

It’s silky black hairs reminded me of a miniature version of the tarantula Pamphobeteus antinous.  It’s broad and prominent coxae reminded me of pictures of large mygalomorphs viewed ventrally.  It also has long pedipalps that almost look like a fifth pair of legs, another trait in common with mygalomorphs.

What stood out to me most was a unique attribute of its life cycle.  Unlike other araneomorphae, and very much like more basal spiders, females continue to molt after reaching maturity, and can live for many years.  This is the reason for one of its many regional names: the southern hibernating spider.  So named because it lives year to year, rather than dying off each winter and hatching in spring as many other spiders do.  

Molt produced by K. hibernalis adult. Photo by C. Peterson

Even today, this particular species captivates me whenever I encounter it.  So today we are going to look at this interesting animal, and try to understand its place in the world of spiders.  

When I was first starting out in the invertebrate hobby, I often found that I had a desire to understand my favorite species in their greater context.  For example, I knew that tarantulas were Mygalomorphs, but I didn’t know what this meant.  I knew that it involved the possession of certain traits, but I didn’t understand why they have the traits they do, or where those features came from.

I found the subject of arthropod taxonomy understandably daunting, full of revisions, huge words that defy pronunciation, and a variety of subjects I would need to familiarize myself with before any of it would make sense.  I suspect this is also the case for many people who enjoy the wonderful world of arthropods.  Over the years, I have grown and done the ground work to open the door to understanding the evolution and classification of arthropods in a general sense.  

For this reason, when I make a post about a particular species or group, I’ll try to include some taxonomic context to help these daunting subjects become a little more manageable, hopefully this will help you appreciate the nature of your favorite animals a little better as well.  

Today We’ll take a quick look at spider taxonomy, which is a huge and complex web (pun intended of course) of relationships that are still being actively researched and refined.   We don’t need to go very deep into the spider family tree, because our subject is very primitive compared to more derived species, it is found near the root of the araneomorphs.

Remember that spiders are arachnids, which are a subset of a group called Chericerates, which are a subset of arthropods emerging during the mid-Cambrian about 500 million years ago.

Approximate higher relationships of major arthropod groups. Illustration by C. Peterson.

As a group, chelicerates represent a more basal arthropod form than, for example, insects and myriapods.  The exact relationships among arthropods is a complex web to untangle, since they are the single most diverse branch of animals there ever was.  Molecular data is being used to establish some revisions, but that is a story for another time.

Spiders represent a subgrouping of arachnids that form the clade Araneae; or simply spiders.  From here, the tree branches again into two groups representing the totality of what are considered to be true spiders; the opisthothelae, which includes virtually all extant spiders, and the mesothelae representing many extinct spiders, and only one extant family; the Liphistiidae.

The name Mesothelae means “middle nipple” named for their spinnerets being placed near the middle of the underside of the abdomen.  Opisthothele means “behind or backward nipple” placing the spinnerets at the back of the abdomen as in most modern spiders. 

Spinneret arrangement relative to hypothetical ancestral state for all spiders. Illustration by C. Peterson

Opisthothelae are further divided into two broad groups; the Mygalomorphae including tarantulas and trap door spiders, and the Araneomorphae which is all the rest.  

Mygalomorph spiders retain a variety of basal spider features including the full set of 4 book lungs, and vertically operating chelicerae, but have lost some other features, most notably the (visibly) segmented abdomen present in most arachnids including basal spiders, and the complete loss of the anterior median pair of spinnerets.  

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Abdominal segmentation typical in arachnids. From top left clockwise: Lephistiid, Amblypygid, Thelyphonid, Scorpion. Images compiled from resources at wikimedia commons, CC.

Most Araneomorphs have lost one set of book lungs. While many retain the anterior pair, they now also feature a modified posterior pair as a trachea.  Their fangs cross each other in a pinching motion, and they retain the anterior median spinnerets, or at least a vestige of it, more on this in a moment. 

The next branch is where our subject can be seen on the horizon.  Traditionally, araneomorph spiders have been split into two main groupings, the entelegynae, which possesses a hardened (or sclerotized) female epigyne and the more basal Haplogynae which lacks these more complex reproductive structures. A third clade, the Hypochilidae, are sister to all other araneomorphs and represent the most basal of all extant araneomorph spiders.

The vast majority of extant spiders are members of Entelegynae.  

Previous high order grouping of spiders. Note that Haplogynae has been found to be paraphyletic and has been revised. See next illustration. Illustration by C. Peterson.

Recently, the grouping Haplogynae has been shown to be paraphyletic, meaning that it included a common ancestor for most (not all) Haplogyne spiders; that is to say that no common ancestor could be identified that included all (and only) Haplogyne spiders. This means that rather than two subdivisions of Araneomorphae, Haplogynae has been split into several groups to better represent the true relationships of extant spiders.  

Likely representation of the relationship of the basal Haplogyene spiders to the rest of Araneomorphae. Note that the group Leptonetidae was previously grouped with the Haplygene spiders, but may represent a sister clade to the Entelegynae. Based on information from this paper. Illustration by C. Peterson.

Molecular studies indicate that haplogynae did include a monophyletic clade, but that a few interesting outliers were lumped in with it, requiring reclassification. Most of the traditional Haplogynae were found to form a clade and are now referred to as Synspermiata, the ecribellate haplogynes.  One family, the Leptonetidae was shown to likely be basal to the entelegynae, though this is tentative. More research will be needed to resolve these relationships in greater detail.

Another family, Filistatidae, was separated from the previous “Haplogynae” and placed as a sister group to Hypochilidae, the so called paleocribellate spiders. This means that Filistatids are among the oldest and most primative of araneomorph lineages. The Hypochilidae-Filistidae group diverged from the rest of the spiders during the mid to late Mesozoic era, 120-294 million years ago. They represent one of the earliest developments in araneomorph history.

The hypochilidae represent the most basal of extant araneomorph spiders.  They possess the defining intersecting fangs, but still have four book lungs like mygalomorphs do.  

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An example of a Hypochilid spider. Hypochilus pococki, image: Kaldari, CC0, via Wikimedia Commons.

Their sister group, the Filistatidae is what we are interested in today.  Defining characteristics of Filistadidae include the retention of the cribellum, a structure derived from the anterior median spinnerets present in primitive spiders.

The cribellum is a silk spinning apparatus present in some araneomorphae, usually more basal clades.  Spiders that retain this feature are referred to as cribellate spiders, those lacking it are said to be ecribellate. Most aranomorphs are ecribellate, but many still retain a visible vestige of this structure called a colulus.

The retention of the cribellum is one of the features that distinguishes the Filistatidae-Hypochilidae clade from Synspermiata, which are a collection of ecribellate Haplogynes.

The cribellum is made of one or more plates with small holes or perforations.  Spiders that retain a cribellum also feature a comb-like row of hairs on the hind legs called a calamistrum. This is used to tease out the non-adhesive silk for the animal to use. The result is a mess of extremely fine silk strands.  This unique type of spider silk is known as cribellate silk.  

Cribellum and spinnerets of adult K. hibernalis. Photo by C. Peterson

Because cribellate silk is not adhesive, the mechanism of prey capture relies on entanglement. The silk traps and entangles prey sort of like Velcro, the more it struggles, the more the tangled and wadded silk stretches and surrounds them.

Some examples of cribellate spiders of other taxons include the ogre faced deinopis, the industrious Agelenopis, a common feature in many backyards, the beautiful Eresus, and the fascinating and enigmatic Uloboridae, notable not only for being cribellate spiders that produce orbs, an advanced form of trapping generally seen in more derived spiders, but they are also entirely nonvenomous, having lost their venom glands at some point in their evolution.

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Examples of other cribellate spiders. Image compiled from resources at wikimedia commons, CC.

With this understanding of the base of the spider family tree in mind, let’s return to our subject today.

Remember that Kukulcania hibernalis is a member of the family filistatidae, a clade of spiders near the base of the araneomorph tree. This spider represents a basal form of what is the beginning of modern spiders. Filistatids are an ancient lineage with their origins in the mid Jurassic, to early Cretaceous period.  

Filistatidae and its sister clade Hypochilidae also do something unique with their cribellate silk; they fold it.  Cribellate silk is already messy but this folding tends to create a very distinctive look for the silken structures of the group.

An example of the characteristically “messy” retreat constructed by Filistatids. Photo on right is K. hibernalis by C. Peterson, Lake Whales Florida, August 2021. Top left: unidentified Filistatid lair, wikimedia commons, CC. Bottom left: Hypochilid web, wikimedia commons CC.

K. hibernalis is one of the most evolutionarily significant spider species you are likely to find in your yard.  Common in dark, dry places like sheds, woodpiles, and underneath porch covers, this species is easily spotted from afar by its distinctively messy webbing.

This species has an enormous distribution, being widely distributed throughout the southeastern United States, Central America, most of South America, and even one confirmed established population in West Africa.

K. hibernalis range. Illustration by C. Peterson.

They tend to prefer areas that are not often disturbed. I located most of the specimens featured in this video essay in an old pump room housing a colony several generations old, in undisturbed wire trays, and among the items on an old shelf. As a child I frequently found them under porches, in old sheds, and around plumbing.

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A collage of different retreat styles exhibited by the animal. When possible, they will create a retreat into a hole or tube and show a preference for vertical or inverted orientation as shown in the top photos. They are resourceful enough to also create retreats made entirely out of silk and debris when no other cover is available, and can use corners to simulate a more secure retreat as demonstrated by the juveniles in the lower images. Photos by C. Peterson, Lake Whales Florida, August 2021.

You can see from the footage/images that they are capable of modifying their retreats depending on the opportunity. Sometimes they create neat holes, sometimes messy conglomerations. They can be oriented vertically, horizontally, or even entirely upside down. Most of the specimens I have encountered seem to display a preference for verticality.

I remember as a child I used to love sneaking out onto my back porch at night and see the large females just outside their silken retreat waiting for food or mates.   These animals are easily startled, retreating at the first sign of danger.  They are also easily teased out with a little movement.

K. hibernalis also displays strong sexual dimorphism common to Filistatids.  Upon maturing, males take on an entirely different color.  Interestingly, the mature male of this species bears a remarkable resemblance to recluse spiders, Loxoceles species. This species is responsible for many false reports of introduced Loxoceles populations due to mistaken identity.

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Comparison of the mature male K. hibernalis (on left) and L. reclusa female (on right). Note that K. hibernalis has a total of 8 eyes clustered closely together similar to mygalomorphs, while Loxoceles species have 6 eyes spread out symmetrically in pairs. More noticeable are the long pedipalps of Kukulcania compared to Loxoceles, and the lack of a “violin” shape on the carapace. Image compiled from resources at wikimedia commons, CC.

As an interesting side note, recluse spiders are part of Synspermiata, being members of the family Sicariidae. Like Filistatids, they were once grouped among the so-called “Haplogyne” spiders, but do not retain a cribellum, and only have six eyes.

Female K. hibernalis continue to molt regularly even after reaching maturity, a trait common to mygalomorphs, but rare among araneomorphae.  

Another interesting feature of the Filistatidae is the temporary appearance of the posterior book lungs in early instars.  An image of the vestigial posterior book lungs taken by scanning electron microscope can be seen on page 150 (figure 97, image B.) in this research paper.

These spiders seem to be remarkably tolerant of each other as well. Adults and juveniles can be found closely clustered together with little evidence of cannibalism. Even large females guarding egg sacs can be found inches away from other spiders.

Some research has been done that suggests that, at least as juveniles, this species could be considered borderline social. See the references below for links to a few papers examining this. Regardless of the degree of social behavior they exhibit, they do seem to be densely concentrated where the do occur, with populations occurring in clusters.

As you might expect, they are easy to keep in captivity, having a low metabolism and low dependence on moisture. A project I am working on is establishing a “colony” of sorts in captivity. I will update new footage and behavior when this project is complete.

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A male approaching a female already guarding an egg sac. Female was nested in the open. Photo by C. Peterson, Lake Whales Florida, August 2021.

One last interesting thing to look at is the mating behavior of this species.  The mating ritual takes a bit of time, the male spends a while announcing his presence, drawing the female out without becoming dinner.  It is also interesting to observe the resemblance to mygalomorph mating.

Overall, Kukulcania hibernalis is an interesting spider.  Its early divergence makes it a classic choice for studies of phylogenics and comparative morphology. This relic of evolution and of my childhood will always catch my attention whenever I encounter it.

So the next time you find a big black spider in a messy web, before you reach for the bug spray, take another look, you may just be encountering something special.

Thank you for reading.  If you enjoy content like this, please consider subscribing to our email list and follow us on YouTube and social media, and be sure to hit the notification bell to be alerted when new content is available.

youtube.com/watch?v=C6LqvevTU1o

Thank you, and see you next time.

Recommended resources:

Paper: “Growth and nest hole size preferences in immature Southern House spiders.” By James Carrel. https://www.researchgate.net/publication/272088007_Growth_and_Nest_Hole_Size_Preferences_in_Immature_Southern_House_Spiders_Araneae_Filistatidae_Are_They_Constrained_Consumers

Paper: “Egg guarding and spiderling group-feeding in crevice weaver spiders” By Kathryn Macdonald and James Cokendolpher. https://www.researchgate.net/publication/228366729_Egg_guarding_and_spiderling_group-feeding_in_crevice_weaver_spiders_Araneae_Filistatidae

Paper: “The crevice Weaver Spider Genus Kukulcania” by Martin Ramirez and Ivan L F Magalhaes. https://www.researchgate.net/publication/331191512_The_Crevice_Weaver_Spider_Genus_Kukulcania_Araneae_Filistatidae

What are clades?

In the last entry, I introduced you to 18th century botanist Carl Linnaeus, and described his major contribution to biology: a universal taxonomic system.   Today, I want to take another look at taxonomy, and discuss some of the flaws in this now dated and obsolete classification system, some solutions to these problems, and what it means for the bigger picture of life on earth.  So join me as we explore life, in endless forms beautiful.  

Carl Linneaus was a swedish botanist who is the author of one of the most versatile and universal systems of taxonomy. His greatest insight was noticing that all creatures fit into a larger pattern of nested fundamental similarities becoming more general the broader one looked, and more specific the narrower one’s focus.

Carl Linnaeus and the basic conception of his classification scheme.

He envisioned a hierarchical system of broadly similar groups gradually diverging into more specific and numerous divisions of organisms culminating with members of reproductively isolated breeding populations called species.  This system of nested hierarchies inadvertently revealed the common ancestry of all life, and was an important contributing factor to the discovery of evolution. 

The linnaean system can be thought of as something like a low resolution snapshot of the history of life.  It worked to group extant organisms into familiar categories, while providing some level of context into how they fit into the larger scheme of biological development.

But for all its strengths, the Linnaean taxonomic system does suffer from a handful of flaws that could not have been predicted in Linn’s day.  For what he had to work with, it was remarkable for its time.  But we now recognize that it is no longer sufficient to contain and describe the vast array of organisms known today.  Let’s take a closer look at why.

First, the Linnaean system assumes equal ranks for all lineages of organisms.  This is because the system does not take the ancestry of life into account, being conceived by creationists who had not yet realized how life develops.  

For example, let’s say we have two lineages of organisms to classify.  

For the first, let’s call it lineage A, we assume a nice simple three level relationship at the bottom which we will call family, genus and species.

An illustration showing that the ancestral relationships between populations may not neatly fall into the predetermined number of ranks conceived in the Linnaean system.

But let’s compare the same level of classification with an organism (B) that has greater diversity in its population.  

We can see that developments within this group, represented by the circles with other groups diverging from them, meaning that there is more between the “genus” and “family” level for this group as opposed to the other one.  

Are these extra divisions at the family level, or the genus level?  The fact that we think to ask this question reveals the problem; because of our previously defined ranks, we assume it must be one or the other.  Put another way, we intuit that life must somehow fit itself into our ranked system, rather than creating our system to take the branching and convoluted nature of life into account.  It is important to note that these ranks are entirely arbitrary categories intended to provide a reference to make it easier to think about populations.

This visual illustrates where the Linnaean system breaks down when there is rapid speciation, large levels of diversity in a lineage, or a lineage that has existed for a long time relative to another, and will therefore have a longer family history.  

Another problem this visual can illustrate is that it fails to accurately reflect evolutionary history.  In this example, we have forced the varying developments into the classic ranks for the sake of consistency.  

An illustration showing how misleading the system becomes when we try to force populations to fit within the scheme.

But how are we to know that the genera are not of equal rank?  Nothing about this arrangement implies the relationship of the grey genera to the dark blue ones, the implication is that they are of equal rank, despite one being the ancestor of the other. 

One way to solve this is by adding additional groups where needed.  Since not all family trees are equal, this eventually causes the Linnaean system to break down into a confusing patchwork with some groups requiring auxiliary categorizations right next to taxons that don’t.  

An illustration showing some of the myriad sub categories inserted into the Linnaean scheme in order to account for extensive layers of diversity. Note that the path through the ranks can be different for different organisms. This undermines the whole premise of ranking.

The problem is that now that we allow as many categories as we need, we quickly run out of Linnaean names. We can add a higher and lower category above and below each rank, but the few extra categories this affords us still does not come close to the number that exist in nature. These inter-ranks are applied inconsistently, due to the fact that different lineages are not symmetrical; thus we are forced to partially disassemble our neat system of equal ranks in order to accommodate varying degrees of relatedness, this renders the system too convoluted to be readily used.

For example, There are over 40,000 species of spiders, who first appeared in the fossil record dating back to just under around 400 million years ago. By contrast, there are 1500 species of rodents, making their first appearance a mere 54 million years ago.  There are about 40 species of cats, appearing 35 million years ago. 

Do you suppose their family trees will be equal in length, and number of branches, and diversity such that they can all be condensed into the same number of equal ranked groupings?

A hypothetical illustration comparing an old and diverse lineage, to a new and less diverse one. Note that older does not always mean more diverse; spiders and scorpions have existed for about the same amount of time (scorpions were likely first), but the diversity of scorpions is only 5% (by species count) that of spiders.

The fact that not all lineages are symmetrical means that the whole idea of predetermined, equal ranks is misguided from the beginning. When we account for all of these additional branches, nodes, forks and long histories, our neat categorical set becomes disconnected and stretched out into something that resembles more of a tree.  

Hypothetical phylogenetic tree demonstrating that predetermined ranks cannot consistently fit the reality of family trees.

A ranked system to describe each member of a family tree for just a few generations of one family would break down with all of the outliers, exceptions and special circumstances that pile up. The more generations we include, the more the pattern of a convoluted tree emerges. This is the crux of it, the Linnaean system is envisioned as hierarchy of linear ranks, but it is attempting to describe a network of interrelated populations spanning the entire range of all life.

This situation is further complicated by the fact that different lineages also have varying degrees of differentiation as well. What this means is that while ancestral relationships are relatively straightforward for exclusively sexually reproductive organisms like animals, the web becomes even more tangled with plants. Many plants are able to hybridize between species, and produce viable offspring such that their lineages mix and match more easily and in more ways than animals.

The only way forward is to simply embrace the branching nature of family trees and take to heart that different lineages are not equivalent, and that the web of life is a lot more convoluted and complex than Linnaeus’ neat and tidy system could ever have accounted for.  This means that the entire concept of an equally ranked system such as Linnaeus initially envisioned is not only impossible, it doesn’t make sense to even try.  

The new taxonomic system uses phylogenetic trees to classify and trace family histories. In this way as many categories as are needed can be made to reflect the evident ancestry.  When we find a lineage of organisms that evidently arose from a common ancestor it forms a distinct branch. When this happens, we say that these organisms form a clade; a group that includes a common ancestor and all its descendants.  Such a clade is monophyletic, because it includes all its descendants and thus represents a single grouping.

An introduction to phylogenetic trees

One benefit is that that these divisions are objectively real, as opposed to arbitrary ranks we can invent for the sake of naming things.  We are able to ask, “do these organisms form a clade?” not “how can we classify them?”  

Here is an example of a monophyletic clade that is familiar to us today.  

An illustration demonstrating monophyletic clades. Note that Aves forms a clade, which is nested within Dinosauria, which is nested beside Crocodilians in the clade Archosauria. This “clades within clades” pattern also demonstrates that an organism cannot outgrow its ancestry, it will always belong to all the clades its ancestors did.

Here is the division of the clade archosauria, one of the two main branches of the reptile tree.  You can see that the division including birds branches from a single point, and contains everything after it.  This means that birds form a clade.  For this reason the clade retains the name of the Linnaean class Aves to denote all birds, also known as avian dinosaurs.  

You may hear a clade be referred to as “unranked.” This is a reference to the Linnaean system that may be more familiar to readers or researchers. What it means is that the clade is not equivalent to any Linnaean rank, and does not exist as a category in Linnaean taxonomy.

You can also see from this cladogram that birds are a subset of the groups making up the clade dinosauria, which makes birds a subset of dinosaurs.  Not all dinosaurs were birds, but all birds are dinosaurs.  

This illustrates another feature of phylogenetic classification, one that it shares in common with Linnaean ranks; every new clade continues to belong to every clade its ancestors did.  Birds are a subset of dinosaurs, dinosaurs are a subset of archosaurs, which are diapsids, which are sauropsids, which are amniotes, which are tetrapods, which are chordates, and so on.  

An illustration demonstrating that all new subsets are always part of the parent set in both systems.

This is because the “nested hierarchies” or “groups within groups” nature that the Linnaean system revealed is a feature of life, and was one of the greatest strengths of the taxonomic system.  One could say that Linnaeus’ innovation was “the grouping of groups” since he did not coin the terms “genus” or “species” but he did have the idea to create and group higher level classifications in such a way so as to smoothly unite all animals into a common category.

Another term this visual can illustrate is what is meant by a “sister clade”.  You can see that Crocodilians are not dinosaurs, but they are archosaurs.  Thus we would say that Crocodilians are a sister clade to dinosaurs.  Pterosaurs are also not dinosaurs, although they are not shown on this diagram.  

Another advantage of using phylogenetic trees for classification that would not have been available during Linneaus’ day is the use of molecular analyses to determine ancestral relationships.  

This is the same kind of test used to determine family histories, or identify the parents of a child, although it is more difficult the more distantly related organisms are. Hence the name, phylogenetics; phylogeny (evolutionary history) based on genetics. 

One final shortcoming of the Linnaean ranks was that many of them were paraphyletic.  This means that the grouping includes a common ancestor, but only some of its descendants. 

A generic representation of the contrast between a monophyletic and paraphyletic taxon. A paraphyletic taxon arbitrarily excludes some of its descendants.

An example of a paraphyletic grouping is the Linnaean class Reptilia. (Although Linnaeus originally lumped reptiles in with amphibians, which only exaggerated this error.)

This would not have been known in Linnaeus day, since he and his colleagues only sought to classify extant species. This resulted in a view of life that had huge “gaps” or “jumps” in it. This is what I meant when I said that this early system was like a snapshot closeup of a small part of a much larger tree.

In this illustration, you can visualize the “low resolution” of the time. The unknown and extinct animals always linked the extant species back to the common ancestor of all animals, but this was a blind spot in Linnaeus day. As our knowledge has increased, we are able to see the relationships between these groups with greater resolution. Turns out there was a lot of information in that blind spot.

An illustration contrasting how increased knowledge of biodiversity has resulted in a better resolved understanding of the history of life. Note that Testudines are not shown on this chart. Their position among the diapsids (last common ancestor of all reptiles and birds) was recently confirmed, but their exact placement is still being actively researched.

The more we learn, the more complete our knowledge the history of life is, and the fewer “gaps” and “jumps” exist among the animal phyla. In the 18th century, we were zoomed in on only the most recent tips of what we now know are distinct branches.

The closer we look, the more details there are between any given creatures, and its route back to the common ancestor of all animals. This chart leaves out vast swaths of creatures, opting to show only the simplest representation of the relationships between these phyla.

Not only are paraphyletic classifications completely arbitrary, they can be misleading, causing us to rank descendant groups as equal to their ancestors.  Linnaean taxonomy provides a catalogue style system for filing different creatures into a database.

But in order to understand the history of life, and the evolutionary context of whatever organism you are looking at, you need a different suite of information. In the Linnaean system, groups may not only fail to intuitively reflect their ancestry, but may actually give the wrong idea entirely. This requires you to bring background knowledge to the table in order to keep track of where you are while using the catalog.

Technically, this causes the Linnaean system to become redundant in additional to being unwieldly. The use of monophyletic clades encodes the evolutionary history into the classification scheme itself such that learning taxonomy is learning phylogeny. This solution has a mathematical elegance.

Those Linnaean ranks that did constitute a clade, like Mammalia and Aves retain their name for the sake of simplicity. These familiar named clades can serve as something like a mile marker to help navigate the vast world of biodiversity, since the number of clades is astronomical.

This is one of the struggles of taxonomy. Biodiversity is so vast that practically infinite numbers of monophyletic clades exist, and are continually produced as populations continue to diverge from each other. Many of these clades are not named, so it helps to start with a familiar landmark, and navigate towards the group we want to look at depending on how specific we are being.

That being said, informal paraphyletic groups can still be useful for specific applications. For example, “fish” is another paraphyletic term, and even though it essentially means nothing in a taxonomic sense, the word is far from useless in the colloquial sense.

Hopefully this essay helped you understand what a monophyletic clade is, where Linnaean taxonomy is misguided, and why paraphyletic taxons can be problematic.  I hope also that I have helped you see how evolutionary history and taxonomy are inseparably linked, as your understanding of one improves, so does the other.

In the next video in this series, I am going to go into more detail of the reptile family tree, expanding it to include extinct animals such as dinosaurs, pterosaurs, ichthyosaurs and evaluate the phylogenetic position of turtles. I hope you’ll join me.