Muscicapa sodhii, J. Berton C. Harris, Pamela C. Rasmussen, Ding Li Yong, Dewi M. Prawiradilaga, Dadang Dwi Putra, Philip D. Round & Frank E. Rheindt, 2014

J. Berton C. Harris, Pamela C. Rasmussen, Ding Li Yong, Dewi M. Prawiradilaga, Dadang Dwi Putra, Philip D. Round & Frank E. Rheindt, 2014, A New Species of Muscicapa Flycatcher from Sulawesi, Indonesia, PLoS ONE 9 (11), pp. 1-33 : 15-29

Muscicapa sodhii , sp. nov.

Sulawesi Streaked Flycatcher

urn:lsid:zoobank.org:act: 54B169C6-CD09-455C-B80E-0FB178611BD4

Holotype

MZB.Ornit.33.525; adult male, collected 23 Jun 2012. Left testis 4×2.5 mm, pale tan in color; no brood patch; no cloacal protuberance (Plate 1, Figure 8). Skull 100% pneumatized; no bursa; no enlarged gape flanges. Little fat; stomach and intestine full of small black items (saved in alcohol). No head, body, wing, or tail molt. Collected by the Prawiradilaga field party; prepared by Pamela C.

Rasmussen, field number LL-PCR-2012-010.

Diagnosis

A small, drab gray-brown muscicapid flycatcher with indistinct facial patterning, strong dusky streaking below, and short primary projection. Differs from

Muscicapa griseisticta in having the bill longer and more strongly hooked but

relatively less broad; a weaker face pattern, with indistinct pale eyering (vs.

prominent), dark spotting on throat (vs. mostly white throat), an ill-defined

malar stripe and no pale moustachial stripe (vs. strong); much shorter and more rounded wing; shorter undertail coverts; and shorter, slightly more notched tail. Differs from Muscicapa sibirica in its longer, deeper, and, in Sino-Himalayan

forms of M. sibirica , broader bill, much weaker head and throat pattern, much clearer streaking below, on a whitish background (vs. mostly dark background, especially in Sino-Himalayan forms); much shorter primary projection and first primary; shorter undertail coverts; and compared to M. s. sibirica , shorter and less notched tail. Differs from all forms of Muscicapa dauurica , as well as M. segregata and M. randi , especially in the strongly streaked underparts of M. sodhii . Differs additionally from M. dauurica in its shorter primary projection (longer only than M. d. umbrosa) and its more strongly hooked and, compared to M. d. dauurica , narrower bill; from M. randi and M. segregata in its shorter tail, less extensive pale area on lower mandible, and longer undertail coverts, and from M. segregata in its narrower bill and shorter tarsi.

The song of the new species is higher-pitched than that of all similar Asian species, and differs from them additionally in its combination of relatively narrow bandwidth, few note repetitions, mostly clear, longer notes, few harmonics, and low similarity between adjacent strophes.

Description of holotype

Color terms using Munsell Color’s [29] notation are shown in Table 10. Bill black, terete, with prominent culmen ridge and fairly strong hook, moderately broad, narrowing evenly toward tip, the distal fifth markedly narrow. Proximal third of lower mandible pale, distinctly so and sharply contrasting with black distal two-thirds of lower mandible, the pale extending around the ramal

branches. Rictal bristles fairly prominent, moderately stiff, black. Forehead graybrown with distinct dark brown centers and pale gray-brown edges, these

markings larger on larger feathers of crown, and becoming obscure on feathers of rear crown. Nape and hindneck nearly uniform gray-brown. Sides of head

uniform gray-brown, as nape. Lores mixed white and gray-brown, the feathers tipped dark giving a grizzled appearance, extends as narrow band over bill and as narrow (one feather row) indistinct eye ring, strongest in front of and below eye, weak above and behind eye. Pale lores grade into face below eye and sides of throat, which are indistinctly speckled. Chin and throat white, lightly and finely speckled gray-brown. There is no marked moustachial or malar streak; there is even gradation from pale grizzled lores to speckled throat and uniform graybrown auriculars.

Speckled throat grades into profusely streaked breast, which grades into more uniform brown breast sides and flanks, and less profusely streaked belly, especially center of belly. Flanks browner, less distinctly streaked than belly sides, and center of lower belly unmarked white. Ground color of entire underparts white. Vent unmarked white. Feathers of lower tibia white.

Mantle as nape. Feathers of back (below mantle) mostly lost during preparation due to shot damage, but same color as mantle. Rump and uppertail coverts plain gray-brown, as mantle. Primaries plain dark brown, darker than body plumage. Tertials paler, especially the outer webs. Secondaries as primaries but with very narrow pale edgings.

Marginal coverts (leading wing edge coverts; [30]) mixed dark brown and white. All upperwing coverts dark gray-brown with indistinct, broad pale grayer edges. Tail uniform drab dark brown, slightly paler and grayer on proximal outer feathers.

Soft part colors (from freshly dead specimen): Iris dark brown, narrow orbital ring black; bill black, basal third of lower mandible pale yellowish-cream; tarsi brownish-black, toepads grayish-tan, claws black.

Measurements of holotype

Body weight 12.5 g; culmen from feathers 10.3 mm, wing length (flattened) 64 mm, tarsus 12.5 mm, tail 44.0 mm.

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Paratype

MZB.Ornit.33.530, unsexed adult (gonads destroyed by pellet), collected 25 Jun 2012 ( Figure 8). No brood patch; no cloacal protuberance. Skull 100% pneumatized; no bursa; no enlarged gape flanges. Little fat; stomach full with several hard, rough, black, 2×2 mm seeds, beetle elytra, and other insect parts (contents saved in alcohol). Relatively large tapeworm found in pieces in abdominal cavity saved in alcohol. No head molt but body molt light, and molt present in primaries, secondaries, and rectrices. Collected by the Prawiradilaga field party; prepared by Pamela C. Rasmussen, field number LL-PCR-2012-014. Body weight 12.0 g; culmen from feathers 11 mm, wing length (flattened) 62 mm, tarsus 13.4 mm, tail 44.5 mm.

Variation in the type series

The holotype is in worn plumage and was not molting, while the paratype was molting but mostly in fresh plumage. Differences in color and pattern are striking in direct comparison—the holotype is much browner above, the paratype much grayer. The holotype lacks distinct pale edges to tertials and secondaries, while the paratype has distinct buff edges to all these, narrow on secondaries and broader on tertials. The holotype has indistinct brown streaking from breast to flanks, with a white center of belly and vent; the throat brown-spotted, most densely on chin and least on lower central throat. The paratype has the same pattern below but the streaks below are bolder, grayer, and broader. The paratype (unlike the holotype) was prepared with one wing partially spread, and its primary underwing coverts are heavily dark streaked, looking mostly dark. The undersides of the primaries and secondaries are dark brown (paler in old feathers) with broad pale tan

edgings, narrowest near tips. Both have similar face patterns, but while there is a narrow pale grizzled area over the bill that meets in the center in the holotype, this Variables deemed most important in bold italic, those of somewhat lesser importance in bold only, and those not considered important in regular font. doi:10.1371/journal.pone.0112657.t009

pale area is interrupted by the bill base in the paratype. The lower face (below plain auriculars) is blotchy, in the holotype with no discernible pale

submoustachial and darker malar area, while in the paratype there is a moderately marked paler submoustachial and dark malar. Mantle feathers when fresh (as in the paratype) look uniform/monotone but actually have broad pale edges/tips on both sides then diffuse dark blotches. When worn (as in the holotype) the pale edges disappear and the feathers are then dark-tipped. Secondary greater coverts are vaguely paler tan-edged in fresh plumage (as in the paratype), not prominently as on secondaries and tertials. In worn plumage (as in the holotype), the wing feathers are much more uniform but the primaries look distinctly darker than the tertials, which wear to medium-pale brown, especially on the lateral webs.

Specimens examined

No other specimens of the new species are known to exist.

Etymology

The new species is named in honor of the late Professor Navjot S. Sodhi (1962– 2011) for his monumental contributions to conservation biology and ornithology in Southeast Asia. Dr. Sodhi played a leading role in elucidating the effects of habitat disturbance on biodiversity, especially birds, across Southeast Asia (e.g. [31–36]). This research provided a fundamental understanding of the ecology of Southeast Asian forests and critical information required for conservation

planning. In addition, Dr. Sodhi was an outstanding mentor for many students including D.L.Y. and J.B.C.H. for whom he was an honours and doctorate supervisor, respectively. The English name recognizes the endemic distribution of the species, with the descriptive term ‘‘Streaked’’ to avoid possible confusion with other endemic Sulawesi flycatchers.

Remarks

Distribution and habitat

The species is known by our two specimens, the sightings and photographs of others, and our song recording from Baku Bakulu, Central Sulawesi. In 1997 King et al. [19] observed the flycatcher in a ‘‘very patchy remnant of forest’’ near Baku Bakulu. During our fieldwork in 2011 and 2012, the Baku Bakulu area was a mosaic of mature cacao plantations and small patches of remnant forest trees ( Figure 9). This mixed agro-forest landscape abuts mature secondary and primary forest in Lore Lindu National Park. Despite the disturbance in the area, we found several forest-dependent species there, including Rufous-throated Flycatcher Ficedula rufigula , Sulawesi Dwarf Kingfisher Ceyx fallax , and Red-bellied Pitta Pitta erythrogaster . The flycatcher was seen at all levels in remnant forest trees in the agro-forest landscape.

There are also several sight records of M. sodhii from Danau Tambing, Badeaha, and the Sedoa area in the Napu Valley on the northeastern side of Lore Lindu National Park. So far, the species is not known from the western side of the park, where fewer birdwatchers visit, but some ornithological studies have been done [35, 37]. Other records in Sulawesi are based on photographs from Matabulu and Toraut, North Sulawesi and Karaenta, South Sulawesi. Despite the number of records to date, the distribution of M. sodhii remains poorly known. Most of the handful of sites where the species has been recorded, like Bantimurung-

Burusaraung and Lore Lindu National Parks, are regularly visited by birdwatchers, and are where the species has been reported multiple times. However, the remote protected areas in Sulawesi’s eastern and southeastern peninsulas are seldom visited and their avifaunas poorly known. The elevational range of Muscicapa sodhii is mainly within an elevational band of 150 m asl to 1,200 m in lowland and submontane evergreen forest and disturbed habitats. There are few records at higher elevations; the occasional observations of the species at Badeaha and Danau Tambing at c. 1,700 m may be exceptional and could involve post-breeding

dispersal. While approximately half of the records are from primary lowland and submontane forests, the frequency of records (including that of the holotype and paratype) in disturbed habitats indicates that the species is tolerant of disturbance. It appears likely, however, that when it occurs in cacao plantations it requires the presence of at least scattered mature native forest trees.

Vocal behavior

According to King et al. [19], the pair of birds they observed at Baku Bakulu gave high-pitched calls, which were recorded and played back, eliciting a noticeable response.

At 1702 h on 21 Jun 2012 at Baku Bakulu, J.B.C.H. found a singing individual of the undescribed flycatcher, sitting inconspicuously at mid-level near a fork of the main trunk of a large tree on a small hill-slope. The bird opened its bill noticeably while singing, and looked around a little, but otherwise sat quite still, making it hard to detect even though it was not hidden by foliage. After making a few recordings, P.C.R. played these quietly back to the flycatcher, which did not exhibit any obvious change in behavior but continued singing until 1707 h. It then flew off and we heard and recorded it singing again until 1722 h in a nearby densely vegetated area, but it did not respond noticeably to playback, nor was it seen again that day. Although differences in song before and after playback were not noted in the field, the later recordings (especially AV#17427–17428) made after playback are composed of short strophes, often of a single motif, while in the earlier-made cuts of the same individual most strophes are considerably longer and contain several motifs.

Behavior

Although the behavior of M. sodhii remains poorly known, scattered

documentation from local and visiting birdwatchers have allowed some aspects of foraging behavior and breeding ecology to be established. Like some other closely related Muscicapa flycatchers, M. sodhii appears to be an obligate insectivore that forages inconspicuously at all levels but perhaps mainly mid-levels. During our field work in 2011 and 2012, we observed single individuals sallying for flying doi:10.1371/journal.pone.0112657.g009 insects, including large damselflies, from perches from several feet above ground to the subcanopy. Other known prey items as inferred from published

photographs include orthopterans, including katydids (www.orientalbirdimages. org). Observations from other birdwatchers (e.g. [38–40]) indicate that the species may sporadically participate in mixed foraging flocks with other small passerines such as whistlers and white-eyes.

We did not observe breeding during our field work in May–Jun, but breeding details have been reported in three birdwatching trip reports. Farrow and Robson [41] and Hutchinson [42] observed adult birds attending single juveniles at Bantimurung-Bulusaraung in early Oct, while Hutchinson [43] reported one adult feeding a juvenile adjacent to Lore Lindu National Park near Wuasa in late Sep. The fact that breeding has only been reported in Sep–Oct despite multiple records (n=17) from every month from Apr to Oct in the Lore Lindu and Bantimurung areas suggests that breeding, at least in central-south Sulawesi, may coincide with the start of the monsoon season. In North Sulawesi, the species is known to breed in May based on photographs deposited in the Oriental Birds Image database (www.orientalbirdimages.org) of a fledgling documented in May 2009.

Diagnosability and species status criteria

Though superficially similar in appearance to M. griseisticta , M. sodhii is a wellmarked species (see Diagnosis) that does not show close morphological approach to any other known taxon. In mtDNA it differs by at least 4.1% from all congeners, and it is highly distinct morphologically from the taxa that it is most similar to in mtDNA. Bearing in mind the small sample of singing individuals and that further vocal sampling is needed, M. sodhii also appears to differ in song from its congeners to a similar and in some cases to a stronger extent than the others do from each other. However, even if the available vocal sample proves

unrepresentative, morphological and genetic differences clearly indicate species status. We therefore consider that, under any modern species concept, M. sodhii unambiguously fits the definition of a species and that its recognition as such will be non-controversial.

Conservation

The fairly wide elevational and distributional range of M. sodhii , and its tolerance of habitat disturbance indicate that it is not immediately at high risk from logging or habitat conversion, which is ongoing in Sulawesi’s lowlands and, increasingly, highlands [17, 18, 44, 45]. The species probably does not occur in young cacao monoculture, or where remnant forest trees are not preserved. It is a low-density species in the Lore Lindu National Park area and it appears to be uncommon elsewhere in Sulawesi, given that the species was only reported in 17 of 51 birdwatching reports we sampled since its first documentation in 1997.

Nonetheless, the paucity of reports may be partly because the species had not yet been formally described. Present knowledge suggests the species does not approach the thresholds for ‘Vulnerable’ under the IUCN’s range size or inferred population trend criteria [46]. We propose that the species be placed in the ‘Least Concern’ category.

Persistence of the species in small forest patches in a mosaic of cacao

plantations over 15 years in Baku Bakulu shows that M. sodhii tolerates some level of disturbance and fragmentation. Our limited data however do not allow us to infer if disturbed habitats are preferred over primary forest. The scientific description of the species, including its voice, should allow comparative surveys to be done in forests and disturbed areas to learn about the species’ habitat preferences. Furthermore, it should be a priority to survey poorly-sampled areas such as Sulawesi’s eastern and southeastern peninsulas to collect distributional information on M. sodhii . Taxonomic studies are also needed to evaluate whether populations on Sulawesi’s northern and southern peninsulas are distinct from Central Sulawesi birds. Many Sulawesi bird genera have racially or specifically distinct representatives that are allopatric on the island’s peninsulas (e.g., Heinrichia , Zosterops , and Ficedula , among others), and it is possible that more than one taxon of Muscicapa is resident on the extraordinarily complex island of Sulawesi.

Tables

Table 6. Plumage scores for six Asian Muscicapa taxa presented as mean ± SD with sample size (n) in parentheses.

M. griseisticta Variable (n = 15) new species (n = 2) M. d. umbrosa (n = 1) M. d. williamsoni (n = 9) M. randi (n = 3) M. segregata (n = 12)
eyering (1= none, 4.3±1.2 10= prominent)	3	4	5.7±1.3	2±1.7	3.1±0.9
crown speckling 3.7±1.3 (1= none, 10= prominent)	6	1	1.8±1.2	3.3±1.2	2.4±1.2
breast color 4.9±0.6 (1= pure white, 10= all brown)	6	9	5.7±1.1	8±1	8.3±0.97
breast streaking 8±0.76 (1= none, 10= heavy)	7	2	6±1.4	2±1	2.5±1.4
vent color 1.1±0.26 (1= pure white, 10= pure brown)	1	1	1.1±0.3	1	1.2±0.39
vent marking 1.1±0.26 (1= none, 10= heavy)	1	1	1	1	1
malar (1= none, 7.5±0.92 10= strong)	1	1	3.2±2.2	1.3 ±0.58	1.4±0.67
rump (1= as mantle, 1 10= much more rufous)	1	3	5.4±1.2	3±1	1.3±0.65
tertials (1= plain, 4±1.9 10= broad strong pale margins)	6±2.8	1	7.6±1.7	5.7±4	4.3±2.1
upperparts darkness 5±0.38 (1= very dark, 10= very pale)	5.5±0.7	3	5.4±0.7	5.3 ±0.58	5.3±0.98
upperparts hue 3.1±0.52 (1= cold, 10= rufous)	2.5±0.7	2	6.8±1.1	3.3±0.58	4.5±1
secondary coverts 3.3±1.3 spotting (1= none, 10= strong)	2	8	6.4±2.4	4.7±4.6	4.4±2.5
lores (1= no pale, 3.8±1.4 10= broad pale)	6±1.4	3	5.8±1.1	5.3±2.3	4.5±1.7
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