A liverwort and hornwort checklist is provided for Peru. The list is based on over 800 literature references, including monographs, regional studies, and molecular investigations. We report 702 accepted species (including 75 species here reported as new to Peru), 41 doubtful species and 60 rejected species previously reported for Peru. This is a substantial increase from the 508 species reported in the previous checklist published in 1984. We validate one name and typify 23 taxa and identify over 100 more names in need of typification. Peru, due to its diverse physical geography and climate, harbours remarkable biodiversity. However, the knowledge of its liverwort flora (and bryoflora in general) remains limited compared to vascular plants, birds, mammals and herpetofauna. We predict that further fieldwork, coupled with scrutiny of collections in Peruvian and other herbaria, will enhance the number of new liverwort records for the country, in particular the addition of regionally widespread species recorded in neighbouring country checklists but not yet from Peru.

BACKGROUND AND AIMS:Like numerous patterns in ecology and evolution, the latitudinal diversity gradient varies across phylogenetic levels. Yet, studies that investigate systematically how patterns and processes change at different phylogenetic levels, from the tips to the root, are still relatively scarce. Here, we test the hypothesis that, despite the high long-distance dispersal capacities of liverworts, which would be expected to result in the homogenization of their distributions, an increase of diversity with latitude persists at increasing phylogenetic level owing to macroclimatic niche conservatism since the earliest evolutionary history of the group. METHODS:Liverwort distributions were scored for 450 operational geographical units worldwide. From the tips to the root, the phylogeny was sliced continuously to examine how taxonomic and phylogenetic diversity are correlated with latitude in a standardized way. Taxonomic diversity and mean phylogenetic distance among taxa were computed for each operational geographical unit at different phylogenetic levels and correlated with macroecological factors using spatial linear models. KEY RESULTS:The correlation between taxonomic diversity and latitude shifted progressively from significantly negative at species level to non-significant, then significantly positive at the highest phylogenetic levels. Taxonomic diversity and mean phylogenetic distance were both significantly correlated with macroclimatic factors across all phylogenetic levels. CONCLUSIONS:In contrast to the marked increase of angiosperm family diversity towards the tropics, the latitudinal diversity gradient evidenced at species level in liverworts decayed progressively at increasing phylogenetic level, suggesting that phylogenetic niche conservatism has played a much weaker role in liverworts than in angiosperms. The inverted latitudinal diversity gradient towards the deepest phylogenetic levels lends support to the hypothesis that the earliest lineages diversified in extra-tropical conditions, explaining why, unlike in angiosperms, high species richness in the tropics is not associated with high phylogenetic diversity in liverworts. Our results highlight the extent to which a phylogenetically deconstructive approach allows for a better understanding of the accumulation of biodiversity through time.

This represents an update to the 2010 checklist of all hornworts and liverworts reported for Java. The checklist includes i) new information published since 2010, ii) previously overlooked historical literature as well as iii) corrected unintentional mistakes that were published in 2010. We now report that the hornwort and liverwort flora of Java consists of 608 known and accepted species as well as several invalid names that we can not place to any taxon. The updated checklist includes 35 recorded species that are considered dubious and another 157 species are excluded from the flora of Java.

The first ever liverwort and hornwort checklist is provided for the Maluku Islands (Moluccas/Spice Islands) of Indonesia. We report 355 accepted and 16 doubtful species and reject 22 species previously reported for Maluku Islands. The list is based on the specimens housed in the Herbarium Bogoriense (BO) and reports from over 500 literature references, including monographs, regional studies, and molecular investigations. The Maluku Islands are part of the Wallacea Biodiversity Hotspot with many unique species found only in Wallacea. Publications focusing on liverworts and hornworts of Maluku Islands are few and scattered. Considering regionally widespread species that have been recorded elsewhere, we predict that further fieldwork exploring the diversity of habitats coupled with collections unveiled from regional herbaria, a number of new records remain to be reported.

Lepidozia parvula N.Kitag. was erroneously treated as a synonym of Lepidozia robusta Steph. in Söderström et al. (2016) due to the unsupported synonymization by Bapna and Kachroo (2000). The taxon has later been recognized (e.g. by Lai et al. 2008; Sukkharak et al. 2008, 2014) and has been reported from a few localities in Thailand. The synonymization must be an error since the two taxa are not at all similar. For instance, Lepidozia parvula has uniseriate leaf lobes (except at the base) while Lepidozia robusta has narrowly triangular leaf lobes. Lepidozia parvula is only known from Thailand while Lepidozia robusta is known from China, Nepal, Bhutan and India, but not from Thailand.

xKochi University, Faculty o f Science and Technology, D epartm ent o f B iological Sciences, 2 -5 -1 A kebono-cho, Kochi, Kochi 780-8520, Japan ; 2Hattori Botanical Laboratory, Obi 6-1-26 , Nichinan City, Miyazaki 889-2535, Japan ; 3Gantz Family Collections Center, The Field Museum, Chicago, IL, USA; 4D epartm ent o f Biology, Norwegian University o f Science and Technology, Trondheim, Norway; *E-mail: lars@ elpt.info

(2969) Colurolejeunea tenuicornis A. Evans in Trans. Connecticut Acad. Arts Sci. 10: 455. Mar 1900 [Hepatics], nom. cons. prop. Lectotypus (hic designatus): Hawaii, Oahu, Kalihi ridge of Nuuanu, 11 Jun 1898, Cooke (YU barcode YU.216401; isolectotypus: G barcode G00115870). (=) Lejeunea ceratophora Nees in Gottsche & al., Syn. Hepat.: 405. 11–13 Dec 1845, nom. rej. prop. Lectotypus (hic designatus): “in cortice peruviano”, hb. Nees. (STR!; isolectotypus: G barcode G00283388). The liverwort Colura tenuicornis (A. Evans) Steph. (Sp. Hepat. 5: 942. 1916), based on Colurolejeunea tenuicornis A. Evans (in Trans. Connecticut Acad. Arts Sci. 10: 455. 1900), is a widespread and well-known pan-tropical taxon also known under several synonyms (see Jovet-Ast in Rev. Bryol. Lichénol. 22: 203–206. 1954). The name is used in many recent publications (e.g., Eggers, Trop. Bryol. 20: 109–115. 2001; He & Zhu in Cryptog. Bryol. 32: 107–112. 2011; Lavocat-Bernard & Schäfer-Verwimp in Cryptog. Bryol. 32: 233–272. 2011; Pócs in Chenia 11: 12. 2013; Schäfer-Verwimp & Van Melick in Cryptog. Bryol. 37: 305–348. 2016; Wigginton in Trop. Bryol. 9: 1–138. 2018; Gradstein in Mem. New York Bot. Gard. 121: 1–723. 2021; Lee & al. in PhytoKeys 199: 29–111. 2022). The C.M. Cooke gathering from Oahu was first indicated as type by Bonner (Index Hepat. 4: 882. 1963) but without identification of the herbarium in which the specimen was preserved. Zhu & So (in Nova Hedwigia Beih. 121: 233. 2001) referred to the specimen at YU as the “holotype”, but this was not an effective lectotypification under Art. 7.11 of the ICN (Turland & al. in Regnum Veg. 159. 2018), lacking the phrase “designated here”. Second-step lectotypification by this specimen is accomplished above. The specimen is to be seen at https://bryophyteportal.org/portal/collections/individual/index.php?occid=2493834). A second specimen, probably detached by Stephani from the type and thus an isolectotype, is in G (G barcode G00115870, to be seen at https://www.ville-ge.ch/musinfo/bd/cjb/chg/adetail.php?id=236382). Lejeunea ceratophora was described by Nees (in Gottsche & al., Syn. Hepat. 3: 405. 1845) from peruvian bark (“cortice peruviano”) (Cinchona officinalis L.), widely assumed to be from Peru, and later reported from a few places in Central America (Standley in Field Mus. Nat. Hist., Bot. Ser. 10: 62. 1931; Herzog in Rev. Bryol. Lichénol. 20: 175. 1952). Jovet-Ast (l.c.) placed it, as Colura ceratophora (Nees) Trevis. (in Mem. Reale Ist. Lombardo Sci., Ser. 3, Cl. Sci. Mat. 4(13): 402. 1877), in synonymy with C. tenuicornis with some doubts, but using the latter, younger name for the taxon, although noting that there was a priority problem. This has been followed since then without anyone proposing to use the older name with the exception of Menzel (in Willdenowia 14: 492. 1984). The type that we have designated, preserved in STR, consists of tiny twig bark pieces with a few shoots in poor condition. A duplicate in G (which we have only seen online at https://www.ville-ge.ch/musinfo/bd/cjb/chg/adetail.php?id=280153) contains a small piece of bark which seems to be in equally bad condition. Search in the online international bibliographic databases JSTOR (http://www.jstor.org), Web of Science (https://apps.webofknowledge.com/), and Google Scholar (http://scholar.google.com/), respectively (using the species names as search terms, accessed 6 Apr 2023), for use of the names in the literature gave the following numbers of hits: Colura ceratophora (1, 0, 1), Colura tenuicornis (29, 4, 174). Colura tenuicornis is the name that has been widely and persistently used for this taxon for almost 70 years. Usage has become well established. In order to maintain a stable nomenclature, we propose conserving Colurolejeunea tenuicornis, the basionym of Colura tenuicornis, against Lejeunea ceratophora, the basionym of Colura ceratophora. LS, https://orcid.org/0000-0002-9315-4978 AH, https://orcid.org/0000-0001-8390-319x TP, https://orcid.org/0000-0002-1359-1298 FB, https://orcid.org/0000-0002-1532-3609 We wish to thank John McNeill and John Wiersema for comments on the proposal and for making us aware of an isotype that we had overlooked.

Five names in Frullania, F. convoluta Lindenb. et Hampe, F. herzogiana Steph. 1911 (= F. rio-janeirensis (Raddi) Ångstr., syn. nov.), F. herzogiana Steph. 1916 (= F. convoluta), F. convoluta var. ampliata Herzog (= F. convoluta, syn. nov.) and F. piliflora var. appendiculata Herzog (= F. peruviana Gottsche, syn. nov.) are lectotypified.

Centralization and standardization of biodiversity data increases accessibility and can lead to the development of checklists and other resources as powerful and important tools for taxonomy and conservation. The publication of new liverwort and hornwort names remains vastly scattered across dozens of journals. Thus we continue the longstanding index series of published names of liverworts and hornworts with 2021 and 2022. The list herein includes the following: six higher taxon names, 15 generic names, 81 infrageneric names, 187 specific names, 32 infraspecific names, three infrageneric autonyms and 13 infraspecific autonyms. Among them are 24 names of fossils as well as six illegitimate and 64 invalid names. Six older names omitted in the earlier indices are included.

Katagiri (2018) synonymized Trichocolea minutifolia and Trichocolea wattsiana under the name Trichocolea hatcheri E.A.Hodgs. overlooking the fact that Trichocolea hatcheri is a younger name. Trichocolea minutifolia and Trichocolea wattsiana were both published in the same paper (Stephani and Watts 1914) and one of them should have been selected. Trichocolea wattsiana has richer type material, so we here select that as the name to be used for this species. For two of the names Katagiri (2018) cited the type as “holotype” although the describing author did not select a holotype and there exists type material in more than one herbarium. As it is after the 1 January 2001, it is too late for the “holotype” of Katagiri (2018) to be corrected to lectotype (cf. ICN Art. 7.10, Turland et al. 2018). Thus, we here correct this and formally designate the same specimens as new lectotypes.

Kuwahara (1984) synonymized Metzgeria madagassa Steph. and Metzgeria thomeensis Steph. under the name Metzgeria australis Steph. So (2004) rejected this synonymy and recognized Metzgeria madagassa from Africa (excluding the syntype from Sikkim) with Metzgeria warnstorffii Steph., Metzgeria camerunensis Steph. and Metzgeria limbatosetosa Steph. as synonyms. Her treatment has generally been followed by subsequent authors of African Metzgeria. However, she overlooked the fact that Metzgeria warnstorffii is the oldest valid name.

Grolle (1989) united Strepsilejeunea subcrenulata Herzog with the younger name Cheilolejeunea papillata Solari. However, transferring the former name to Cheilolejeunea was blocked by the existence of the earlier name Cheilolejeunea subcrenulata (Spruce) R.M.Schust. so Grolle used next available name which was Solari's Cheilolejeunea papillata. Unfortunately, the latter name is also a later homonym to Cheilolejeunea papillata (Herzog) Kachroo, which Grolle overlooked. Thus, the taxon is lacking a legitimate name in Cheilolejeunea. Therefore, we here replace the earliest name with a new name in the genus.

Twenty-two names of liverwort and hornwort taxa described from Central America are lectotypified. Most of the taxa have been identified as types earlier but not formally selected. Two taxa are supplementarily lectotypified as the original lectotypification could be applied on more than one specimen.

Lepidozia novae-zelandiae var. heterostipa R.M.Schust. ex J.J.Engel, var. nov. Based on Lepidozia novae-zelandiae var. heterostipa R.M.Schust., Fieldiana, Bot. (n.ser.) 42: 70, 2001 (Engel and Schuster 2001), nom. inval. (ICN2018 Art. 40.7; herbarium not specified). Holotype: New Zealand, North Is., Tongariro Natl. Park, Tree Trunk Gorge, along west bank of Tongariro River, 15 Feb. 1995, Schuster 95-772 (F-C0550810F https://fm-digitalassets.fieldmuseum.org/1939/387/C0550810F.jpg).

A widely accessible authoritative list of known plant species is a fundamental requirement for local, regional, and global biodiversity conservation efforts. The longstanding index series of published names of liverworts and hornworts continues with 2019 and 2020. The list herein includes one higher taxon name, six generic names, 13 infrageneric names, 124 specific names, 27 infraspecific names, one infrageneric autonym and nine infraspecific autonyms for 2019 and 2020. The list includes 13 names of fossils as well as 13 invalid names. Four older names omitted in the earlier indices are included.