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Honeycutt2009chap76.Pdf Rodents (Rodentia) Rodney L. Honeycutt to either features of the zygomasseteric system (con- Division of Natural Science, Pepperdine University, 24255 Pacifi c A guration of the infraorbital foramen and placement Coast Highway Malibu, CA 90263-4321, USA (rodney.honeycutt@ of the masseter muscles in the jaw) or the angle of the pepperdine.edu) lower jaw relative to the plane of the incisors (15, 16). Tullberg’s classiA cation (15) identiA es two suborders, Abstract Sciurognathi and Hystricognathi, based on the angle of the lower jaw, and the hystricognathous condition sup- The Order Rodentia is the most diverse group of mammals, ports the monophyly of a clade containing phiomorph represented by 34 extant families. Recent molecular data and caviomorph rodents (Fig. 2). Nevertheless, the have helped to clarify relationships among these families, phylogenetic distribution of features of the infraorbital and provide a framework for the higher-level classifi ca- foramen reveals evidence of parallel changes within tion of rodents. Molecular time estimates among rodent Rodentia (17–19). families have been made in several studies, permitting a Detailed molecular phylogenetic studies contrib- timetree of rodent evolution to be constructed. The time- ute greatly to the resolution of the rodent evolutionary tree shows a partitioning of rodent families into three major tree, and areas of congruence among the majority of clades and reveals three periods of diversifi cation: The late these studies provide strong support for many interfa- Cretaceous, 88–66 million years ago (Ma), the Paleocene milial relationships among rodents (Fig. 2). For instance, to early Eocene, 60–55 Ma, and the late Oligocene to early two major monophyletic groups, Hystricomorpha and a Miocene, 25–15 Ma. squirrel-like clade, are supported by nuclear (2, 19–22) 7 e Order Rodentia (represented by squirrels, mice, rats, guinea pigs, and others) contains 42% (2277) of all spe- cies and 39% (481) of all genera described for the Class Mammalia (1) (Fig. 1). If one includes the recently dis- covered extant Family Diatomyidae (2), the order con- tains 34 families (1). All early classiA cations of mammals (3) and more recent cladistic analysis of morphological variation (craniodental, postcranial, fetal membrane) (4) support a monophyletic Rodentia. Although several early molecular phylogenetic studies questioned the validity of a monophyletic Rodentia (5, 6), more recent molecu- lar data, based on either nuclear DNA (7–9) or a more thorough analysis of sequences of whole mitochondrial genomes (10), reveal support in favor of monophyly. In this review I will address phylogenetic relationships and divergence times for families within Rodentia. Even though the assignment of species and genera of rodents to speciA c families is well established (1), the derivation of a family-level phylogeny for rodents is more challenging (11–14). 7 is problem has resulted in a variety of classiA cations that vary in terms of recog- nized suborders and the morphological features used to deA ne monophyletic groups (1, 11, 15–17). Two of the Fig. 1 A capybara (Hydrochaeris hydrochaeris), Family Caviidae. major characteristics used to diagnose suborders relate Credit: R. L. Honeycutt. R. Honeycutt. Rodents (Rodentia). Pp. 490–494 in e Timetree of Life, S. B. Hedges and S. Kumar, Eds. (Oxford University Press, 2009). HHedges.indbedges.indb 449090 11/28/2009/28/2009 11:30:10:30:10 PPMM Eukaryota; Metazoa; Vertebrata; Mammalia; Rodentia 491 Dasyproctidae 20 Agoutidae 22 18 Caviidae Erethizontidae 16 Dinomyidae 27 Chinchillidae 17 Abrocomidae Octodontidae Caviomorpha 26 28 Ctenomyidae 13 25 Capromyidae 29 Echimyidae 30 Hystricomorpha Myocastoridae 11 Bathyergidae 15 Petromuridae 21 7 Thryonomyidae Phiomorpha Hystricidae Diatomyidae 12 Ctenodactylidae Anomaluridae 2 10 Peditidae 4 Dipodidae 6 Spalacidae 14 Nesomyidae 3 23 Cricetidae 24 1 Mouse-like Muridae Castoridae 5 Geomyidae 19 Heteromyidae Gliridae 8 Sciuridae 9 Aplodontidae Squirrel-like Squirrel-like Late K Paleogene Neogene MESOZOIC CENOZOIC 75 50 25 0 Million years ago Fig. 2 A timetree of rodents (Rodentia). Divergence times are shown in Table 1. The muroid Families Calomyscidae (no divergence time data) and Platcanthomyidae (recently extinct) were not included. Abbreviation: K (Cretaceous). and mitochondrial (18, 24, 25) sequences, and some clades are somewhat more tenuous, but many molecu- nuclear markers (2, 7, 21), including the presence/ lar studies place the squirrel-like clade at the base of absence of retrotransposon insertion loci (9), support a the phylogeny (7, 8, 20, 21, 23). Recent molecular stud- third mouse-like clade. Relationships among these three ies also help resolve the placement of several historically HHedges.indbedges.indb 449191 11/28/2009/28/2009 11:30:13:30:13 PPMM 492 THE TIMETREE OF LIFE Table 1. Divergence times (Ma) and their confi dence/credibility intervals (CI) among rodents (Rodentia). Timetree Estimates Node Time Ref. (2) Ref. (21) Ref. (24) Ref. (27) Ref. (28) Ref. (30) Ref. (31) Time CI Time CI Time Time CI Time CI Time CI Time 192.9––96.9105–85– –––––– – 282.872.180–6493.5106–77– –––––– – 378.967.876–6090101–68– –––––– – 476.066.575–5885.4101–65– –––––– – 566.463.372–5584.993–6650.9–––––– – 666.065.473–578782–5345.7–––––– – 764.261.363–5483.797–6747.6–––––– – 862.457.365–5078.890–6251.2–––––– – 951.550.859–4365.382–5338.3–––––– – 1048.356.865–49–– 39.7–––––– – 1147.845.451–4043.151–29– 5563–46––––– 1244.344.351–38–– – – –––– – 13 41.7 42.7 47–37 38.1 54–16 – 49 54–43 – – 36.6 39–34 – 1439.0–––– – –––––– 39 15 38.0 38 43–33 – – – 45 48–41 – – 30.5 34–27 – 16 35.3 34.3 37–30 – – – – – 37.7 40–35 33.8 36–32 – 1732.233.336–2932.847–2430.4–––––– – 1831.731.735–27–– – –––––– – 19 28.4 27.3 33–22 – – – 27 29–25 – – 31.4 33–29 – 20 26.0 – – – – – 24 26–21 – – 27.9 30–26 – 2125.824.543–20–– – 2728–25––––– 2225.624.128–20–– – ––––26.529–24– 2325.5–––– – ––––––25.5 2424.2–––– – ––––––24.2 25 21.3 18.6 22–15 – – – – – 27.4 32–23 17.5 20–15 – 2621.0–––– – ––––20.623–18– 2720.221.426–18–– – ––––19.122–16– 2817.3–––– – ––19.623–171517–13– 2910.712.415–10–– – 910–7––––– 30 7.3 – – – – – 6 7–5 – – 8.6 10–7 – Note: Node times in the timetree represent the mean of time estimates from different studies. In ref. (30), confi dence interval is based on estimates of standard deviations provided. In ref. (27), divergence time represents an average of two estimates of the same gene (VWF) derived from nucleotide and amino acid sequences, and the CI refers to the range of these estimates. In ref. (21), divergence time represents an average of the GHR and BRCA1 genes; only values derived from the rate-smoothing method were used; CI represents the combine rate of values for both genes. In ref. (28), divergence time represents an average of estimates derived from nuclear (GHR) and mitochondrial (12S rRNA) combined genes, and CI is the range of values. In ref. (31), estimates for Node 14 based on GHR gene, optimized with rate smoothing, and dates for Nodes 23 and 24 represented values from a concatenation of all genes and optimization with the penalized likelihood method. problematic taxa, including the Families Pedetidae Tullberg’s (15) Suborder Hystricognathi, character- (Springhaas), Anomaluridae (scaly-tailed squirrel), ized by a hystricomorphous zygomasseteric system and Geo myidae/Heteromyidae (pocket gopher and pocket a hystricognathous lower jaw, is strongly supported mouse), and Castoridae (beaver) (2, 7, 9, 21, 24). by molecular studies (2, 9, 18, 20, 21, 25 ). In addition, HHedges.indbedges.indb 449292 11/28/2009/28/2009 11:30:13:30:13 PPMM Eukaryota; Metazoa; Vertebrata; Mammalia; Rodentia 493 monophyly of the South American Caviomorpha (guinea 7 e timetree does provide a framework for testing pigs and relatives) is strongly supported, suggesting a hypotheses related to the distribution and diversiA ca- single invasion from African ancestors, represented tion of rodents. 7 e phylogeny in combination with today by the Phiomorpha (2, 7, 9, 20, 26). 7 e placement estimates of divergence times (Fig. 2 and Table 1) reveals of Old World porcupines (Hystricidae) is more contro- three major diversiA cations of rodents, one occurring in versial, yet the majority of data, based on nuclear and the late Cretaceous and early Paleocene, another in the mitochondrial sequences, suggests a basal position for Paleocene to early Eocene, and another that involved the family (2, 20, 21, 23, 27). Relationships among fam- the diversiA cation of African phiomorphs and South ilies within the Caviomorpha are well resolved with a American caviomorphs in the late Oligocene to early combination of nuclear and mitochondrial sequences, Miocene. 7 is pattern is similar to a recent proposal for and with few exceptions (e.g., clade containing Chin- mammalian radiations that suggests an early origin for chilidae and Dinomyidea, Fig. 2) these results are simi- many lineages followed by an increase in diversiA cation lar to groups previously deA ned based on morphology during the Eocene and Oligocene (36). Many rodent fam- (22, 23, 25, 27–29). 7 e current classiA cation of rodents ilies in the timetree also reveal long terminal branches assigns the 34 families to A ve suborders (1), and molecu- and shorter internodes, features that make the resolution lar data support the monophyly of these A ve suborders: of the phylogeny more challenging. Sciuromorpha, Castorimorpha, Myomorpha, Anoma- 7 e timetree also provides insight into the biogeog- luromorpha, and Hystricomorpha (Fig. 2). 7 ree of these raphy of hystricognath rodents. For instance, cavio- suborders comprise the mouse-like clade. As indicated morph rodents A rst appear in the fossil record of South earlier, placement of some of these major groups (e.g., America 37–31 Ma (39) and the oldest phiomorphs in basal squirrel-like clade and the clade uniting Muroidea, Africa date to 37–34 Ma (38), and these dates are simi- Dipodidae, Peditidae, and Anomaluridae, Fig.2) is not lar to those indicated in Table 1.
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