Suncus

Savi's pygmy shrew Suncus etruscus, one of the smallest living mammals, photographed by Daniel Heuclin.

Belongs within: Soricidae.

Suncus, the musk shrews, are found in southern Asia and Africa. One species, the house shrew Suncus murinus, has become widely dispersed through the Indo-Pacific region as a commensal of humans.

Characters (from Nowak 1999): Head and body length 35-150 mm, tail length 25-100 mm. Colour grey to brown. Fur short, soft, may be velvety. Tail with long hairs projecting beyond short bristly hairs. Males with strong musky odor produced from large well-developed scent gland on flanks.

<==Suncus
    |--S. ater IT07
    |--S. dayi IT07
    |--S. etruscus B78
    |--S. fellowesgordoni B78
    |--S. hosei IT07
    |--S. infinitesimus [incl. S. chriseos] B78
    |--S. lixus IT07
    |--S. madagascariensis IT07
    |--S. malayanus IT07
    |--S. mertensi IT07
    |--S. montanus IT07
    |--S. murinus (Linné 1766) K92
    |    |--S. m. murinus B74
    |    `--S. m. tytleri B74
    |--S. remyi IT07
    |--S. stoliczkanus IT07
    |--S. varilla [incl. S. orangiae] B78
    `--S. zeylanicus IT07

*Type species of generic name indicated

REFERENCES

[B74] Bugge, J. 1974. The cephalic arterial system in insectivores, primates, rodents and lagomorphs, with special reference to the systematic classification. Acta Anatomica 87 (Suppl 62): 1-160.

[B78] Butler, P. M. 1978. Insectivora and Chiroptera. In Evolution of African Mammals (V. J. Maglio & H. B. S. Cooke, eds) pp. 56-68. Harvard University Press: Cambridge (Massachusetts).

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PLoS One 2 (3): e296.

[K92] Klompen, J. S. H. 1992. Phylogenetic relationships in the mite family Sarcoptidae (Acari: Astigmata). Miscellaneous Publications, Museum of Zoology, University of Michigan 180: i-vi, 1-154.

Nowak, R. M. 1999. Walker's Mammals of the World vol. 2. JHU Press.

Dormaaliinae

Partial cranium and jaws of Macrocranion nitens, from Novacek et al. (1985).

Belongs within: Lipotyphla.

The Dormaaliinae are an Eocene group of small mammals of uncertain relationships. Novacek (1985) regarded them as belonging to the Erinaceomorpha, among the relatives of the hedgehogs. Butler (1988) pointed out that the dormaaliine Macrocranion lacked the synapomorphies of Lipotyphla and excluded it from that group, but did not propose a specific alternative position. One subgroup, the amphilemurines, had bunodont molars superficially similar to those of Primates.

Characters (from Novacek 1985): P_1-2 reduced, single-rooted; P₃ very small, single-rooted; P₄ premolariform with large protoconid, small metaconid and paraconid, short weakly basined or unbasined talonid with one or two minute cuspules; M_1-3 with paraconids transverse, sharp, crestiform, closely approximated to metaconids; trigonids on M_1-3 distinctly canted anteriorly, anteroposteriorly compressed in occlusal view; P⁴ with sweeping metastylar crest, vestigial or absent metacone, weak posterolingual extension of cingulum; M^1-2 with well-developed hypocone.

<==Dormaaliinae
    |--Dormaalius N85
    |--Macrocranion nitens B88
    `--Amphilemurini N85
         |--Pholidocercus N85
         |--Amphilemur Heller 1935 N85, S62
         |    `--A. eocaenicus Heller 1935 S62
         |--Alsaticopithecus Hürzeler 1947 N85, S62
         |    `--*A. leemanni Hürzeler 1947 S62
         `--Gesneropithex Hürzeler 1946 N85, S62
              `--G. peyeri Hürzeler 1946 S62

*Type species of generic name indicated

==References==

[B88] Butler, P. M. 1988. Phylogeny of the insectivores. In The Phylogeny and Classification of the Tetrapods, vol. 2. Mammals (M. J. Benton, ed.) pp. 117-141. Clarendon Press: Oxford.

[N85] Novacek, M. J. 1985. The Sespedectinae, a new subfamily of hedgehog-like insectivores. American Museum Novitates 2822: 1-24.

Novacek, M. J., T. M. Bown & D. Schankler. 1985. On the classification of the early Tertiary Erinaceomorpha (Insectivora, Mammalia). American Museum Novitates 2813: 1-22.

[S62] Simons, E. L. 1962. A new Eocene primate genus, Cantius, and a revision of some allied European lemuroids. Bulletin of the British Museum (Natural History): Geology 7 (1): 1-36.

Erinaceidae

West European hedgehog Erinaceus europaeus, photographed by Hrald.

Belongs within: Erinaceomorpha.

The Erinaceidae include the hedgehogs and their spine-less cousins, the gymnures. Erinaceids are first recorded from the Middle Eocene, with Entomolestes from North America belonging to the Tupaiodontinae. The Oligocene saw the appearance of the other subfamilies, with Proterix in North America and the remainder (including the tupaiodontine genera Tupaiodon and Ictopidium) in Eurasia. Representatives of the living subfamilies Erinaceinae and Galericinae, as well as the Brachyericinae, reached Africa and North America during the Miocene (Butler 1988).

Characters (from Butler 1988): P⁴/M₁ shear; P⁴ with high paracone and long metacrista; M₁ enlarged with extended trigonid, paraconid forwardly placed as high paracristid. Posterior molars reduced; paraconid on M_2-3 reduced to transverse crest at level of talonid of preceding tooth; M₂ metacrista very short. Zygomatic process of maxilla shifted forward.

<==Erinaceidae
    |--Tupaiodontinae B88
    |    |--Tupaiodon Matthew & Granger 1924 B88, N85
    |    |    |--T. minutus B88
    |    |    `--T. morrisi B88
    |    |--Ictopidium B88
    |    |    |--I. lechei Zdansky 1930 GH99
    |    |    `--I. tatalgolensis B88
    |    |--Entomolestes B88
    |    |    |--E. grangeri B88
    |    |    |--E. nitens M68
    |    |    `--E. siegfriedti M68
    |    `--Zaraalestes GH99
    `--+--Proterix Matthew 1903 B88, G68
       |    |--*P. loomisi Matthew 1903 G68
       |    `--P. bicuspis (Macdonald 1951) [=Apternodus bicuspis] G68
       `--+--Neurogymnurus G68 [Neurogymnurinae B88]
          |--Galericinae B88
          |    |--Tetracus B88
          |    |--Lanthanotherium N85
          |    `--Galerix [incl. Pseudogalerix] N85
          |         |--G. africanus Butler 1956 Bu78
          |         |--G. exilis G68
          |         `--G. socialis N85
          `--+--Brachyericinae [Brachyericini] B88
             |    |--Brachyerix G68
             |    |--Metechinus G68
             |    |--Dimylechinus G68
             |    `--Exallerix B88
             `--Erinaceinae B88
                  |--Protechinus salis Lavocat 1961 Bu78
                  |--Mioechinus oeningensis Bu78
                  |--Postpalerinaceus vireti Crusafont & Villalta 1948 Bu78
                  |--Paraechinus micropus Bu78, V02
                  |--Palaeoscaptor N85
                  |--Amphechinus G68 [incl. Palaeoerinaceus Bu78]
                  |    |--A. edwardsi M60
                  |    |--A. kansuensis M60
                  |    |--A. minimus M60
                  |    `--A. rusingensis Butler 1956 Bu78
                  |--Gymnurechinus Bu78
                  |    |--G. camptolophus Butler 1956 Bu78
                  |    |--G. leakeyi Butler 1956 Bu78
                  |    `--G. songhorensis Butler 1956 Bu78
                  |--Hemiechinus Bu78
                  |    |--H. aethiopicus IT07
                  |    |--H. auritus V02
                  |    |--H. collaris IT07
                  |    |--H. hypomelas IT07
                  |    |--H. megalotis GM71
                  |    |--H. micropus IT07
                  |    `--H. nudiventris IT07
                  `--Erinaceus Linnaeus 1758 P04
                       |  i. s.: E. amurensis IT07
                       |         E. concolor IT07
                       |         E. europaeus Linnaeus 1758 K92
                       |         E. frontalis Bi78 [=Atelerix frontalis IT07]
                       |         E. sansaniensis Bu78
                       `--E. (Atelerix) Bu78
                            |--E. (A.) albiventris Wagner 1841 Bu78, K92
                            |--E. (A.) algirus Bu78
                            |--E. (A.) broomi (Broom 1937) [incl. Atelerix major Broom 1937] Bu78
                            `--E. (A.) sclateri Bu78

Erinaceidae incertae sedis:
  Echinosorex AM02 [Echinosoricinae Bu78]
    `--E. gymnurus AM02
  Hylomys M58
    |--H. hainanensis IT07
    |--H. sinensis IT07
    `--H. suillus M94
  Podogymnura IT07
    |--P. aureospinula IT07
    `--P. truei IT07
  Mesechinus IT07
    |--M. dauuricus IT07
    `--M. hughi IT07
  Neotetracus M58
  Meterix latidens M60

*Type species of generic name indicated

REFERENCES

[AM02] Asher, R. J., M. C. McKenna, R. J. Emry, A. R. Tabrum & D. G. Kron. 2002. Morphology and relationships of Apternodus and other extinct, zalambdodont, placental mammals. Bulletin of the American Museum of Natural History 273: 1-117.

[Bi78] Bigalke, R. C. 1978. Present-day mammals of Africa. In Evolution of African Mammals (V. J. Maglio & H. B. S. Cooke, eds) pp. 1-16. Harvard University Press: Cambridge (Massachusetts).

[Bu78] Butler, P. M. 1978. Insectivora and Chiroptera. In Evolution of African Mammals (V. J. Maglio & H. B. S. Cooke, eds) pp. 56-68. Harvard University Press: Cambridge (Massachusetts).

[B88] Butler, P. M. 1988. Phylogeny of the insectivores. In The Phylogeny and Classification of the Tetrapods, vol. 2. Mammals (M. J. Benton, ed.) pp. 117-141. Clarendon Press: Oxford.

[G68] Gawne, C. E. 1968. The genus Proterix (Insectivora, Erinaceidae) of the Upper Oligocene of North America. American Museum Novitates 2315: 1-26.

[GH99] Gheerbrant, E. & J.-L. Hartenberger. 1999. Nouveau mammifère insectivore (?Lipotyphla, ?Erinaceomorpha) de l’Eocène inférieur de Chambi (Tunisie). Paläontologische Zeitschrift 73 (1-2): 143-156.

[GM71] Goodman, M., & G. W. Moore. 1971. Immunodiffusion systematics of the primates I. The Catarrhini. Systematic Zoology 20 (1): 19-62.

[IT07] Isaac, N. J. B., S. T. Turvey, B. Collen, C. Waterman & J. E. M. Baillie. 2007. Mammals on the EDGE: conservation priorities based on threat and phylogeny. PLoS One 2 (3): e296.

[K92] Klompen, J. S. H. 1992. Phylogenetic relationships in the mite family Sarcoptidae (Acari: Astigmata). Miscellaneous Publications, Museum of Zoology, University of Michigan 180: i-vi, 1-154.

[M94] MacPhee, R. D. E. 1994. Morphology, adaptations, and relationships of Plesiorycteropus, and a diagnosis of a new order of eutherian mammals. Bulletin of the American Museum of Natural History 220: 1-214.

[M58] McDowell, S. B., Jr. 1958. The Greater Antillean insectivores. Bulletin of the American Museum of Natural History 115 (3): 113-214.

[M60] McKenna, M. C. 1960. The Geolabidinae: a new subfamily of early Cenozoic erinaceoid insectivores. University of California Publications in Geological Sciences 37 (2): 131-164.

[M68] McKenna, M. C. 1968. Leptacodon, an American Paleocene nyctithere (Mammalia, Insectivora). American Museum Novitates 2317: 1-12.

[N85] Novacek, M. J. 1985. The Sespedectinae, a new subfamily of hedgehog-like insectivores. American Museum Novitates 2822: 1-24.

[P04] Popov, V. V. 2004. Pliocene small mammals (Mammalia, Lipotyphla, Chiroptera, Lagomorpha, Rodentia) from Muselievo (north Bulgaria). Geodiversitas 26 (3): 403-491.

[V02] Vyas, R. 2002. Pale hedgehog Paraechinus micropus in southeastern Rajasthan. Journal of the Bombay Natural History Society 99 (3): 501.

Erinaceomorpha

Lower jaw of Sespedectes stocki, from Novacek (1985).

Belongs within: Lipotyphla.
Contains: Erinaceidae.

The Erinaceomorpha includes the modern hedgehogs (Erinaceidae) and their fossil relatives. Most fossil species are only known from their dentition, which is relatively unspecialised, and the inclusion or otherwise of taxa in this group has often been debated. According to Butler (1988), the earliest erinaceomorphs include Adunator from the Palaeocene of Europe and North America, with Adapisorex and Leipsanolestes also Palaeocene. Most other non-erinaceid erinaceomorphs are Eocene in age, though Ankylodon is Oligocene.

Characters (from Rose 2006): Dental formula 3.1.4.3/3.1.4.3; molars low-crowned, bunodont; M^1-2 with moderately to well-developed hypocones; fourth premolars premolariform to submolariform.

<==Erinaceomorpha [Erinaceoidea]
    |  i. s.: Neohylomys hainanensis BP87
    |         Talpavoides N85
    |         Opisthopsalis V66
    |         Hyracolestes V66
    |         Praolestes V66
    |         Chambilestes Gheerbrant & Hartenberger 1999 [Chambilestidae] GH99
    |           `--*C. foussanensis Gheerbrant & Hartenberger 1999 GH99
    |--+--Neomatronella N85
    |  `--Adapisorex [Adapisoricidae] B88
    |       `--A. abundans B88
    |--+--Diacodon Cope 1875 B88, C76
    |  |    |--D. alticuspis Cope 1875 C76
    |  |    |--D. celatus Cope 1875 C76
    |  |    `--D. minutus S35
    |  `--Adunator [incl. Diacocherus, Mckennatherium] B88
    |       |--A. ladae B88
    |       |--A. lehmani B88
    |       `--A. minutus B88
    `--+--Leipsanolestes siegfriedti Simpson 1928 B88, S35 [=Leptacodon (Leipsanolestes) siegfriedti S35]
       |--Erinaceidae B88
       `--+--Dartonius B88
          `--+--Talpavus B88
             `--Scenopagidae B88
                  |--Scenopaginae B88
                  |    |--Scenopagus McKenna & Simpson 1959 M60
                  |    |    |--S. curtidens GH99
                  |    |    |--S. edenensis B88
                  |    |    `--S. mcgrewi M60
                  |    |--Ankylodon B88
                  |    |    |--A. annectens Patterson & McGrew 1937 M60
                  |    |    `--A. progressus Galbreath 1953 M60
                  |    `--Embassis Cope 1873 M60
                  |         `--*E. alternans Cope 1873 M60
                  `--Sespedectinae B88
                       |--Crypholestes Novacek 1976 N85
                       |    `--*C. vaughni Novacek 1976 N85
                       `--+--Proterixoides Stock 1935 N85
                          |    `--*P. davisi Stock 1935 N85
                          `--Sespedectes Stock 1935 N85
                               |--*S. singularis Stock 1935 N85
                               `--S. stocki Novacek 1985 N85

*Type species of generic name indicated

REFERENCES

[BP87] Burton, J. A., & B. Pearson. 1987. Collins Guide to the Rare Mammals of the World. Collins: London.

[B88] Butler, P. M. 1988. Phylogeny of the insectivores. In The Phylogeny and Classification of the Tetrapods, vol. 2. Mammals (M. J. Benton, ed.) pp. 117-141. Clarendon Press: Oxford.

[C76] Cope, E. D. 1876. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4 (2): i-iv, 1-370.

[GH99] Gheerbrant, E. & J.-L. Hartenberger. 1999. Nouveau mammifère insectivore (?Lipotyphla, ?Erinaceomorpha) de l’Eocène inférieur de Chambi (Tunisie). Paläontologische Zeitschrift 73 (1-2): 143-156.

[M60] McKenna, M. C. 1960. The Geolabidinae: a new subfamily of early Cenozoic erinaceoid insectivores. University of California Publications in Geological Sciences 37 (2): 131-164.

[N85] Novacek, M. J. 1985. The Sespedectinae, a new subfamily of hedgehog-like insectivores. American Museum Novitates 2822: 1-24.

Rose, K. D. 2006. The Beginning of the Age of Mammals. JHU Press.

[S35] Simpson, G. G. 1935. The Tiffany fauna, Upper Paleocene. I.–Multituberculata, Marsupialia, Insectivora, and ?Chiroptera. American Museum Novitates 795: 1-19.

[V66] Van Valen, L. 1966. Deltatheridia, a new order of mammals. Bulletin of the American Museum of Natural History 132 (1): 1-126.