After the dinos, tree shrews came down and got big!

During the reign of the dinosaurs
tree shrews, like Ptilocercus (Fig. 1) and Tupaia (Fig. 1), stayed in the trees, evolving into tree-dwelling members of the Carnivora (Genetta, Fig. 1), Volitantia (bats, pangolins and dermopterans), Glires (including multituberculates led by Tupaia) and Primates (Microcebus, Fig. 1) in the large reptile tree (LRT, 1818+ taxa) distinct from all gene studies and all other prior trait studies (due to taxon exclusion). The LRT is the first study that found tree-dwelling Caluromys (Fig. 1), an extant tree shrew-like marsupial, as the proximal outgroup to the Placentalia. Based on chronological bracketing, Caluromys relatives lived in the Early Jurassic.

Figure 1. Mammals at the base of the Placentalia include the outgroup taxon: Caluromys, a basal placental: Genetta, a basal Carnivora: Eupleres, a basal Volitantia: Ptilocercus, a basal Primates: Microcebus, and basal Glires: Tupaia.

Figure 1. Mammals at the base of the Placentalia include the outgroup taxon: Caluromys, a basal placental: Genetta, a basal Carnivora: Eupleres, a basal Volitantia: Ptilocercus, a basal Primates: Microcebus, and basal Glires: Tupaia.

After the Cretaceous some tree shrews became terrestrial.
Leptictids, elephant shrews (Rhynchocyon, Fig. 2 and tenrecs (Tenrec) were phylogenetically among the first of the former tree shrews to become fully terrestrial. They were all small. After the Cretaceous some terrestrial tree shrew descendants began to increase in size. Some became elephants, others horses, still others baleen whales, all following Cope’s Rule.

Figure 7. Rhynchocyon, a living elephant shrew, is a living leptictid.

Figure 2. Rhynchocyon, a living elephant shrew, is a living leptictid and a former tree shrew.

Once established on the ground
and spreading beyond the jungles, the following Early Paleocene terrestrial placentals became cat to tiger size: Onychodectes (Fig. 3), Alcidedorbignya (Fig. 3) and Pantolambda (Fig. 3).

Figure 3. Onychodectes, Alcidedorbignya and Pantolambda are former tree shrews now terrestrial of increasing size in the Early Paleocene.

Figure 3. Onychodectes, Alcidedorbignya and Pantolambda are former tree shrews now terrestrial of increasing size in the Early Paleocene. Note the lost of sharp claws replaced by pre-hooves.

By the late Paleocene
taxa like massive Barylambda showed further increases in size. This taxon was basal to giant glyptodonts and ground sloths, some of which ultimately became smaller and returned to the trees as tree sloths.

Figure 1. Barylambda looks like a large ground sloth for good reason. It is a sister to the direct ancestor and nests at the base of the Xenarthra along with Orycteropus, the aardvark.

Figure 4. Late Paleocene Barylambda looks like a large ground sloth for good reason. It is a sister to the direct ancestor and nests at the base of the Xenarthra along with Orycteropus, the aardvark.

PS… saving the best for last.
Writing this blogpost inevitably brought my gaze back to Fruitafossor (Luo and Wible 2005), a small, Late Jurassic digging mammal with four robust fingers, xenarthran lumbars and single cusp, tubular teeth. When first encountered and based on these traits the LRT mistakenly nested Fruitafossor with edentates for the last four years. That Late Jurassic temporal discontinuity in an otherwise Tertiary clade of edentates required a review and revision of taxon scores for Fuitafossor. That review ultimately re-nested Fruitafossor more plausibly and parimoniously basal to echidnas in the LRT. Fruitafossor is a basal echidna from Colorado. That story comes to you tomorrow.


References
Luo Z-X and Wible JR 2005. A late Jurassic digging mammal and early mammal diversification. Science 308:103–107.

 

 

Why hedgehogs and moonrats are not good outgroup taxa for tree shrews

Li and Ni 2016 put together a few Oligocene teeth
and mandibles and introduced us to tiny Ptilocercus kylin (Fig. 1), a sister to the extant treeshrew, Ptilocercus lowii (Fig. 1). Unfortunately, too little is known of the new species to add it to the large reptile tree (LRT, 1786+ taxa, subset Fig. 4). The posterior premolar is the largest tooth in the mandible in both species. I could not find that trait elsewhere among tested taxa.

Figure 1. Several specimens of Ptilocercus kylin (Oligocene) matched to Ptilocercus lowii (extant).

Figure 1. Several specimens of Ptilocercus kylin (Oligocene) matched to Ptilocercus lowii (extant).

Li and Ni 2016 provided a cladogram
(Fig. 2) in which they ‘selected’ (their word, not mine) a moonrat (Echinosorex) and hedgehog (Erinaceus) as outgroup taxa to the tree shrews and their proximal allies.

Figure 2. Cladogram from Li and Ni 2016 showing their inappropriate selection for an outgroup, the hedgehog, Erinaceus.

Figure 2. Cladogram from Li and Ni 2016 showing their inappropriate selection for an outgroup, the hedgehog, Erinaceus.

Li and Ni 2016 report,
“The phylogenetic analysis was based on a data matrix derived from published data in ref. 12 
[= Ni et al. 2013]. In total, 1857 characters were scored for 177 taxa. The 1857 characters comprise 485 dental, 202 cranial, 309 postcranial, 203 soft tissue characters and 658 molecular characters. Two erinaceid insectivores, Erinaceus europaeus and Echinosorex gymnura, were selected as outgroup taxa. The ingroup comprises 13 treeshrews, 28 flying lemurs and plesiadapiforms, 45 lemuriforms and adapiforms, 40 tarsiiforms and 49 anthropoids. Of these taxa, 47 are extant species.”

Never ‘select’ = ‘cherry-pick’ outgroup taxa. Always let your wide gamut cladogram tell you which taxa are outgroups for your more focused studies.

Unfortunately, Li and Ni chose derived taxa, a hedgehog and a moonrat, both members of the gnawing clade, Glires, as their outgroup clade. In the LRT (subset Fig. 4) all members of Glires are derived from the tree shrew, Tupaia. So hedgehogs and moonrats are not basal, primitive outgroups to all tree shrews. Just the opposite. Tree shrews are among their ancestors.

That mistake puts a small crimp on any results Li and Ni obtain. In the LRT basal members of the clade Carnivora would be more suitable as outgroup taxa. Or, better yet, they could have taken it back one or two more nodes to the outgroup taxon for the clade Placentalia, Caluromys (Fig. 3). This extant arboreal wooly opossum, looks and acts more like a placental tree shrew…with a pouch.

Figure 1. Mammals at the base of the Placentalia include the outgroup taxon: Caluromys, a basal placental: Genetta, a basal Carnivora: Eupleres, a basal Volitantia: Ptilocercus, a basal Primates: Microcebus, and basal Glires: Tupaia.

Figure 3. Mammals at the base of the Placentalia include the outgroup taxon: Caluromys, a basal placental: Genetta, a basal Carnivora: Eupleres, a basal Volitantia: Ptilocercus, a basal Primates: Microcebus, and basal Glires: Tupaia. See the LRT for node placement.

Li and Ni 2016 report,
“Approximately 20 hours were required to finish the unconstrained search. More than 769 billion rearrangements were examined. Forty-seven trees with a best score of 13067 were retained. Nearly 21 hours were required to finish the backbone-constrained search. More than 728 billion rearrangements were examined. Twenty trees with a best score of 13141 were retained. The best trees and the strict consensus of these trees were described in PAUP.

The Li and Ni 2016 cladogram would have been better served by deleting all molecular data and adding more basal placental taxa, as in the LRT (Fig. 4). Dental traits are subject to reversals and convergences. Try to use the whole skeleton and use fewer dental traits, as in the LRT. Usually just a few minutes are required to get nearly full resolution of subsets within the LRT.

Figure 3. Subset of the LRT focusing on the clade Scandentia (tree shrews) and the three arboreal clades that arise from it.

Figure 4. Subset of the LRT focusing on the clade Scandentia (tree shrews) and the three arboreal clades that arise from it. See the LRT for more basal taxa.

Li and Ni report,
“We define the Order Scandentia as a crown group, including the clade stemming from the most common ancestor of extant ptilocercids and tupaiids.”

This definition for Scandentia = Placentalia sans Carnivora + basal Placentalia (in the LRT). So it works! This definition is monophyletic in the LRT. Of course, that means the order Scandentia also includes odontocetes, hedgehogs, rats, multituberculates, elephants, humans, giraffes and ground sloths, etc. etc…. everything, but Carnivora and some arboreal post-marsupial civets. Clearly that was not their intention. So… what do we do with this?


References
Li Q and Ni X 2016. An early Oligocene fossil demonstrates treeshrews are slowly evolving “living fossils”. Sci. Rep. 6, 18627; doi: 10.1038/srep18627 (2016).
Ni X, Gebo DL, Dagosto M, Meng J, Tafforeau P, Flynn JJ and Beard KC 2013.The oldest known primate skeleton and early haplorhine evolution. Nature 498, 60–64. This paper is about Archicebus achilles, a tarsier, we looked at earlier here.

 

The basalmost primate in the LRT is alive and living in Madagascar!

Now you have a choice.
Either go out looking for crumbling bits and pieces of basal primate jaws and teeth over vast stretches of badlands… Or go to Madagascar to study basal primates in the wild, and have them feeding from your hand, according to the latest addition to the LRT.

The gray mouse lemur,
(Microcebus murinus; Figs. 1, 2) nests at the base of the all the tested primates in the large reptile tree (LRT, 1692+ taxa; subset Fig. 3), basal to both larger adapid lemurs, Notharctus and Smilodectes.

Figure 1. The gray mouse lemur (Microcebus murinus) nests basal to primates in the LRT.

Figure 1. The gray mouse lemur (Microcebus murinus) nests basal to primates in the LRT.

This largest species in this smallest genus of primates
also nests between two tree shrew taxa, Tupaia (basal to Glires) and Ptilocercus (Fig. 4; basal to Volitantia).

Though living today in Madagascar forests,
Microcebus likely radiated during the Cretaceous, prior to the splitting of Madagascar from Africa 88 mya. Later it gave rise to all extinct and extant adapids and lemurs on that island.

Millions of years ago lemurs were
worldwide in distribution. Now only a few lemurs find refuge in Madagacar. and only in Madagascar.

Figure 2. The skull of Microcebus murinus from Digimorph.org and used with permission. Here colors mark bones.

Figure 2. The skull of Microcebus murinus from Digimorph.org and used with permission. Here colors mark bones.

Microcebus murinus (Miller 1777) is the extant gray mouse lemur an omnivore found only in Madagascar. This nocturnal arboreal basalmost primate in the LRT forages alone, but sleeps in groups, sharing tree holes during the day. Twin babies are typical. Offspring can reproduce after one year. Lifespan extends to ten years. The eyes are large, typical of nocturnal mammals. Relatives include Hapalodectes and Ptilocercus. Descendants include Notharctus and Smilodectes.

The newly expanded clade Scandentia (tree shrews) now unites
Volitantia (bats + pangolins + colugos), Primates and Glires (rodents, rabbits, multituberculates and kin) in the LRT, subset Fig. 3). The addition of Microcebus as the smallest lemur held the possibility that it was the most basal form or one leading to smaller galagos and tarsiers. This time Microcebus turned out to be more primitive.

Figure 3. Subset of the LRT focusing on the clade Scandentia (tree shrews) and the three arboreal clades that arise from it.

Figure 3. Subset of the LRT focusing on the clade Scandentia (tree shrews) and the three arboreal clades that arise from it.

With the addition of Microcebus to the LRT,
the extant pen-tailed tree shrew, Ptilocercus (Fig. 4) nests basal to colugos, which also lack upper incisors. That means an older, more plesiomorphic fossil taxon with a complete set of upper incisors is out there waiting to be discovered somewhere in Early Jurassic fossil beds.

Figure 4. Ptilocercus is a sister to Microcebus nesting with colugos.

Figure 4. Ptilocercus is a sister to Microcebus nesting with colugos.

Paleontologists have been looking for the ancestor of primates,
colugos and bats for ages. They find fewer and smaller bony scraps the deeper they look.

Here’s a solution:
Add extant taxa. Phylogenetic analyses that includes extant taxa can sometimes help by nesting late survivors at basal nodes. Sure the fossil taxa are the real ancestors. Sure, living lemurs are late survivors, radiating into new morphologies and niches, but the soft, cuddly, active chatterboxes (Fig. 1) are still worth studying and scoring.


References
Miller JF 1777. Cimelia Physica p.25

wiki/Microcebus

Shrew opossums (caenolestids) are supposed to be marsupials

According to Wikipedia,
“The family Caenolestidae contains the seven surviving species of shrew opossum: small, shrew-like marsupials that are confined to the Andes mountains of South America.”

Figure 1. Caenolestes skull and in vivo.

Figure 1. Caenolestes skull and in vivo. It sure looks more like a shrew than an opossum. Skull images from Digimorph.org and used with permission. Colors added.

The trouble is
tested caenolestids, Caenolestes (Fig. 1) and Rhyncholestes (Fig. 2), do not have a pouch. Nor do they nest with marsupials in the large reptile tree (LRT, 1412 taxa, subset Fig. 3). But female caenolestids do have a marsupial-like double vagina (see below).

On the traditional side,
Dr. Darren Naish reported online for Tetrapod Zoology/Scientific American in 2015, “Incidentally, the most frequently used name for the group – shrew-opossums – might not be a particularly good one, seeing as they don’t look much like shrews, don’t live like shrews, and don’t act like shrews. And they’re not technically opossums, either, but perhaps we can let that go.”

Contra Dr. Naish’s amusing musings,
shrew opossums nest with placental shrews alongside the otherwise extinct Apatamys (Fig. 3) + Trogosus (Fig. 4) in the Glires clade. All are derived from a tree shrew taxon close to Tupaia. It’s unfortunate that Dr. Naish did not test these taxa while they were on his mind in 2015. That’s how initial errors become perpetuated as long-standing traditions.

Figure 1. Skull of Rhyncholestes along with in vivo photo.

Figure 2. Skull of Rhyncholestes along with in vivo photo.

Rhyncholestes raphanurus (Osgood, 1924; long-nosed shrew-opossum, Chilean shrew opossum, extant; snout-vent length 20cm), nests in the large reptile tree between the shrew-mole, Uropsilus, and the tree shrew, Tupaia at the base of the Apatemys clade. all within the placental clade, Glires. Wikipedia and other sources consider this shrew-like South American mammal a marsupial, but Wiki also notes that Rhyncholestes lacks a marsupium (pouch).

Figure 2. Apatemys nests as a proximal sister to bats in the Halliday et al. tree. But it shares very few traits with bats. Note the very odd dentition.

Figure 3. Apatemys nests as a proximal sister to bats in the Halliday et al. tree. But it shares very few traits with bats. Note the shrew-opposum/rodent-like dentition.

Genetically
Wikipedia reports. “Genetic studies indicate that they are the second most basal order of marsupials, after the didelphimorphs” (Nilsson et al. 2010). That’s exactly where the LRT documents the splitting of eutherian mammals from the phytometatherians and carnimetatherians.  Even so, we’re talking about deep time here. Don’t trust genes. Test traits.

Figure 3. Subset of the LRT focusing on primates and basal glires, including the caenolestids, Caenolestes and Rhyncholestes.

Figure 4. Subset of the LRT focusing on primates and basal glires, including the caenolestids, Caenolestes and Rhyncholestes.

According to AnimalDiversity.org, “In general, members of family Caenolestidae can be distinguished from other marsupial groups by their unique dentition. Their lower middle incisors are large and have a forward slope; likewise, they have a reduced number of incisors. The dental formula for genus Caenolestes is: I 4/3, C 1/1, P 3/3, M 4/4, 46 teeth total. Shrew opossums have short robust limbs, each containing 5 digits; their middle 3 digits are shorter than the outside two. Their humeri are extremely heavy; in comparison, their femurs are relatively slender. Members of family Caenolestidae have unusual lip flaps, they may function as a method of preventing debris from interfering with their whiskers or they may help prevent ingestion of unwanted debris. Similar to other marsupials, Caenolestid females have 2 uteri and 2 vaginas. Members of genus Caenolestes lack a pouch but do have 4 mammae, 2 on either side of their abdomen.”

Unfortunately
the LRT tests only skeletal material, not for ‘number of uteri and vaginas’. While Larry Martin and Darren Naish might wave this trait about in support of a marsupial affinity, the LRT documents the emergence of placentals from marsupials. So the reappearance of a long-lost trait, like a long tail, a sixth digit, or double vaginas is well within the realm of possibilities in placentals.

As a matter of fact,
a double vagina sometimes occurs in humans.

Here, as elsewhere in paleontology,
maximum parsimony is the only yardstick. PAUP is free to nest taxa wherever 231 unbiased scores indicate it should. Moving the two caenolestids to the Metatheria adds 12 steps to the MPT.

The Apatamyidae is a clade that was long considered extinct.
Now it joins several other clades that are no longer extinct, thanks to the LRT.

Rhyncholestes raphanurus (Osgood, 1924; long-nosed shrew-opossum, Chilean shrew opossum, extant; snout-vent length 20cm), nests in the LRT between the shrew-mole, Uropsilus, and a large living shrew, Scutisorex, all within the placental clade, Glires. Wikipedia and other sources consider this shrew-like South American mammal a marsupial, but Wiki also notes that Rhyncholestes lacks a marsupium (pouch).

Caenolestes fuliginosus (originally Hyracodon fuliginosus Tomes 1863)

Apatemys chardini (Marsh 1872, Eocene, 50-33 mya) was a squirrel-lke arboreal herbivore with a massive skull. Here it nests with Trogosus and Tupaia, a tree shrew. It had long slender fingers, a long flexible lumbar region, and a long gracile tail.


References
Marsh OC 1872. Preliminary description of new Tertiary mammals. Part II. American Journal of Science 4(21):202-224.
Nilsson MA, et al. (6 co-authors) 2010. Tracking Marsupial Evolution Using Archaic Genomic Retroposon Insertions”. PLoS Biology. 8 (7): e1000436. doi:10.1371/journal.pbio.1000436
Osgood WH 1924. Field Mus. Nat. Hist. Publ., Zool. Ser. 14:170.

tetrapod-zoology/you-never-hear-much-about-shrew-opossums/
wiki/Shrew_opossum = Caenolestidae
animaldiversity.org/accounts/Caenolestes_fuliginosus/
wiki/Apatemyidae
wiki/Rhyncholestes
wiki/Caenolestes
wiki/Paucituberculata
wiki/Uterus_didelphys

Click here for Glires skulls compared.

Apatemys revisited with DGS

Another short one today
in which the skull elements of Apatemys chardini (Marsh 1872, Eocene, Figs. 1, 2) are restored to their in vivo positions as determined by molar occlusion and jaw glenoid insertion.

Figure 1. Apatemys skull traced and reconstructed using color overlays (DGS). Yes, quite a bit of the mandible appears to be hidden beneath the broken coronoid process.

Figure 1. Apatemys skull traced and reconstructed using color overlays (DGS).

Apatemys chardini (Marsh 1872, Eocene, 50-33 mya) was a squirrel-lke arboreal herbivore with a massive skull. Here it nests between the much larger Trogosus and the more plesiomorphic, Tupaia, a tree shrew. Apatemys had long slender fingers, a long flexible lumbar region, and a long gracile tail.

This taxon also gives rise to the shrew Scutisorex (check out the similar teeth, for instance), and the former tenrecs, Limnogale and Potamogale. All three are extant.

Figure 1. Apatemys, only complete fossil skeleton of an apatemyid, turns out to be a basal shrew. So this clade is not extinct.

Figure 2. Apatemys, only complete fossil skeleton of an apatemyid, turns out to be a squirrel-like  basal shrew. So this clade is not extinct.

References
Marsh OC 1872. Preliminary description of new Tertiary mammals. Part II. American Journal of Science 4(21):202-224.

wiki/Apatemyidae

 

Rodent, rabbit, tree shrew and multituberculate skulls compared

In an effort to understand 
a clade that was giving me trouble on the LRT, I put together the following set of skulls (Fig. 1, click here to enlarge) from the redefined clade Glires (still rodents and rabbits, but also their closest kin).

Figure 1. Click to enlarge. See text for explanation.

Figure 1. Click to enlarge. See text for explanation.

In the above illustration.
Select members of the Glires (rodents, rabbits and relatives, all derived, ultimately from the basal placental, Monodelphis, not to scale. Numbers refer to columns:

  1. Tupaia is a tree shrew. Macroscelides is an elephant shrew. Chrysochloris is a golden mole.
  2. Scutisorex is a hero shrew. Apatemys is a an arboreal apatemyid. Trogosus is a terrestrial apatemyid or tillodont. 
  3. Solenodon is a solendontid. Zalambdalestes is another solenodontid. 
  4. Nambaroo is a primitive rabbit and a kangaroo mimic. Brachyerix is an extinct hedgehog. Gomphos is an extinct rabbit. Orytolagus is an extant rabbit.
  5. Carpolestes is an arboreal plesiadapiform. Plesiadapis is another arboreal plasiadapiform. Taeniolabis is a multituberculate. 
  6. Shenshou is a pre-rodent. Paramys is an extinct rodent. Rattus is an extant rodent. Ignacius has not been tested but usually nestes with plesiadapiform. 
  7. Kryptobaatar is a multituberculate. Ptilodus is a multituberculate. Rugosodon is a multituberculate. Megconus is another multituberculate often considered a mammaliaform.

Still have not found evidence
that multituberculates nested in a clade more primitive than placentals. You’ll note that Zalambdalestes now nests with Solenodon despite the epipubes (found occasionally in other placentals, too). Glires nests between Carnivora and Ptilocercia. See yesterday’s post on basal placentals for basalmost taxa.

 

The golden mole is an underground tree shrew!

Updated October 11, 2016 with a new lateral view of the Chrysochloris skull. 

One more nail in the ‘Afrotheria’ coffin here.
The Cape golden mole (Chrysochloris asiatica, Linneaus 1758; 8-20cm in length) nests with the elephant shrew (Macroscelides) and together they nest with the tree shrew (Tupaia) in the large reptile tree, which scores morphology, not molecules. Prior studies nested golden moles with elephant shrews, tenrecs, aardvarks, hyraxes, elephants and sea cows in the purported clade ‘Afrotheria.’ The large reptile tree does not confirm such a clade.

Figure 1. The Golden Mole (Chrysocloris asiaticus) nests with the tree shrew and elephant shrew in the large reptile tree, not the common mole. Image copyright Digimorph.org and used with permission.

Figure 1. The Golden Mole (Chrysocloris asiaticus) nests with the tree shrew and elephant shrew in the large reptile tree, not the common mole. Image copyright Digimorph.org and used with permission.

Convergent with
Talpa, a true or common mole, and marsupial moles, the golden mole has taken to a life underground, with reduced ears and eyes, large fore claws and enormous shoulder muscles. Note the displacement of the scapulae to the top of the neck. And the pisiform evolves to become a second ulna! (Fig. 2).

Figure 2. Golden mole forelimb. The elongate pisiform (amber) essentially creates a second ulna. Only digits 1-4 are present.

Figure 2. Golden mole forelimb. The elongate pisiform (amber) essentially creates a second ulna. Only digits 1-4 are present. Image copyright Digimorph.org and used with permission.

Rather than broad claws
the golden mole third ungual is large and sharp. The other digits are reduced and digit 5 is absent. The lateral temporal fenestra and the jaw adductors therein, have been squeezed out as the squamosal laminates to the cranium (Fig. 3). And if you look close enough, there is a postorbital bar. But it too, is laminated to the cranium. The posterior jugal is tall and robust.

Figure 3. The lateral view of the golden mole (Chrysochloris) skull shows the tiny orbit and the complete  circumorbital bones, the jugal and frontal.   Image copyright Digimorph.org and used with permission.

Figure 3. The lateral view of the golden mole (Chrysochloris) skull shows the tiny orbit and the complete circumorbital bones, the jugal and frontal.
Image copyright Digimorph.org and used with permission.

 Like the tenrecs,
golden moles possess a cloaca (combined anus and genitals), and males lack a scrotum, but they are not as closely related to tenrecs. Like the Marcoscelides and Tupaia the tibia is longer than the femur in Chrysochloris, which is odd for a tetrapod that does not sprint.

Figure 2. Macroscelides proboscideus, the elephant shrew or sengis is NOT more closely related to elephants with the purported 'Afrotheria.' but instead is related to Tupaia, the tree shrew.

Figure 2. Macroscelides proboscideus, the elephant shrew or sengis is NOT more closely related to elephants with the purported ‘Afrotheria.’ but instead is related to Tupaia, the tree shrew.

Analysis indicates that certain tree shrews
in the Cretaceous descended to the ground. Elephant shrews became sprinters to defend themselves from dinosaur predators. Golden moles burrowed.

References
Linnaeus C 1758. Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.

wiki/Golden_mole
wiki/Cape_golden_mole