﻿Licariaramiroi (Lauraceae), a new species from Western Mexico

﻿Abstract Licariaramiroi, a species endemic to western Mexico, is described and illustrated. The ascription of the new species to Licaria is analysed. It is classified in the subgen. Licaria and is most closely related to L.triandra and L.siphonantha from which it differs by its glabrous vegetative and floral structures, stamens mainly with free anthers and the inner wall of the receptacle sericeous. According to the IUCN criteria, the species is classified as critically endangered.

ics of America and Asia (van der Werff and Richter 1996;van der Werff 2009b). The Lauraceae include 120 species in southern Mexico, which are considered to represent between 90 and 93% of the Mexican Lauraceae (Lorea-Hernández 2002).
Licaria Aubl. is a Neotropical genus of around 60 species, of which at least 20 have been described in the last 25 years (van der Werff 2009a). Of the total number of Licaria species, 17 have been recorded for southern Mexico, eight of which are considered endemic to the country (Lorea-Hernández 2002). Species of Licaria are easily recognised by the presence of fertile stamens in whorl III only, bisporangiate anthers and fruits on a well-developed cupule, usually with a double margin (Burger and van der Werff 1990;van der Werff 1991;Lorea-Hernández 1999;Kurz 2000;van der Werff 2009a). The subgenera Armeniaca, Canella and Licaria are recognised by the anther shape and direction of opening and the presence or absence of staminodes in series I and II, with the latter of the three subgen. presenting the greatest species richness and distribution ranges from Florida on the Atlantic side and Mexico (Sinaloa) on the Pacific slope, to southern Brazil and Bolivia (Burger and van der Werff 1990;Kurz 2000;van der Werff 2009a).
As part of a gradient analysis study of the vegetation of the El Tecolote canyon in the Sierra de Manantlán, Jalisco, Mexico (Cuevas 2002), material was collected from a Licaria that could not be identified. Initially, only sterile material or material with fruits was available, then material with flowers was collected. However, given the characteristics observed in the flower, particularly the variation in the number of stamens, it was decided to wait until more collections with flowers became available. After making detailed studies of the material from the Sierra de Manantlán, we found no accommodation amongst the known species of Licaria, for that reason, this is proposed as a new species.

Materials and methods
Species morphology was studied and described, based on specimens available in the ZEA Herbarium of the University of Guadalajara and on fresh material collected in the field. Vegetative and reproductive structures were carefully revised under a stereoscopic microscope using the protocol recommended by Radford et al. (1974) in order to elaborate the description. Relevant literature was reviewed, including taxonomic treatments of the genus Licaria and descriptions of new species in the genus from the last 25 years (Burger and van der Werff 1990;van der Werff 1991;Gómez-Laurito and Cascante 1999;Lorea-Hernández 1999;Kurz 2000;van der Werff and Vicentini 2000;Lorea-Hernández 2005;van der Werff 2009a). Photographs of type specimens of species that could be related to the Licaria species, proposed here as new, were reviewed in the Tropicos (2022) (www.tropicos.org) and JSTOR (2020) (http://plants. jstor.org) databases. Measurements and photographs of the floral structures were taken using fresh or rehydrated material; all other structures were measured in dehydrated specimens. Microphotographs were taken with a SteREO Zeiss Discovery V12 stereoscopic microscope connected to an AxioCam 305 Color Zeiss camera and Zen 3.2 (blue edition) software. Photographs of the species in the field were taken with a Canon EOS M3 camera. Habitat and phenology data of the new species were obtained from herbarium specimen labels and fieldwork, while information pertaining to the related species was taken from the relevant literature (Kurz 2000;Lorea-Hernández 2005).
Distribution and ecology. Licaria ramiroi is known from only one locality in the Sierra de Manantlán, Jalisco, Mexico. It has been recorded between 1000 and 1600 m in elevation in an ecotone between subdeciduous tropical forest and montane cloud forest. Dominant trees in this habitat correspond to individuals of the Lauraceae family, such as Beilschmiedia manantlanensis Cuevas  Field observations were carried out for three years; as we found no flowers or fruits during the different seasons of the year, nor observed any seedlings, in this period, this leads us to suggest that this species could have supra-annual flowering and fruiting periods.
The locality from where the new species of Licaria is described is an area that has been prominent for records of new species, such as Beilschmiedia manantlanensis (Cuevas & Cochrane, 1999), Sideroxylon brucebenzii (Cuevas & Vázquez-García, 2021) or disjunct distributions, such as those of Desmopsis trunciflora (Schltdl. & Cham.) G.E. Schatz  and Nectandra rudis C.K. Allen (Cuevas et al. 2008), amongst others. Fortunately, the area forms part of the Sierra de Manantlán Biosphere Reserve, ensuring a certain degree of conservation of the flora of this area, which is notable in the region for its uniqueness and the distribution of some of its species (Cuevas 2002).
The species has been recorded in flower in the months of July-August and with ripe fruits from February to March.
Etymology. The species honours the memory of Ramiro Cuevas Guzmán, dear brother of the first author, a person who loved the countryside and who collected plants for more than 30 years together with the author of the species, in various regions of the southern coast of Jalisco. Several of these collections are deposited in national herbaria and some have resulted in scientific novelties.
Preliminary conservation status. According to the Categories and Criteria of the IUCN Red List (IUCN 2012), L. ramiroi is assigned a preliminary status of "Critically Endangered" CR B2a C2a(ii). The known and estimated geographical distribution of this species is less than 10 km 2 in area and it has been recorded at only one locality, with a population size estimated at fewer than 250 mature individuals. Moreover, it should be noted that there is little natural regeneration of the species and few juvenile individuals present. Notes. Due to its erect tepals, the exserted stamens with their filaments fused at the base, the anthers with large locules orientated in the longitudinal axis of the flower, with the valves opening upwards, Licaria ramiroi is classified in the subgen. Licaria, within the group of species with exserted stamens and with the locules on the external surface of the anthers (Kurz 2000). Morphological characters with narrow leaves with 12 or more secondary veins, the shape and size of the flowers, exserted stamens and the large sporangia covering most of the anther body and running lengthwise to the flower axis, relate Licaria ramiroi to L. siphonantha Lorea-Hern. but separates from this species by its fused tepals at its base, generally shorter and glabrous inflorescences, free anthers and the hypanthium internally sericeous (see Table 1). The shape and size of the flowers, the tepals fused at their bases, the glands often fused at their bases with free apices, the size of the cupules in the fruit with strongly undulating margins, as well as the shape and size of the fruit, relate L. ramiroi with the variable L. triandra (sensu Kurz (2000), including L. cervantesii (Kunth) Kosterm. and Misanteca nayaritensis Lundell), the only species of Licaria recorded for western Mexico (Kurz 2000), from which it is separated by the presence of stamens fused only in the filaments, while the anthers are generally free, an internally sericeous hypanthium and other characters that are presented in Table 1. A review of 60 flowers from two individuals (20 from one and 40 from the other) collected 23 years apart, suggests that the proportion of flowers with 3, 4 and 5 stamens in whorl III is more or less equal. This phenomenon observed in L. ramiroi is something previously not recorded in the genus Licaria. The evidence supporting the notion that they are all stamens of whorl III is that all of the anthers are extrorse and there is incomplete separation of stamen primordia and, when this occurs, some of these stamens can show a certain degree of fusion at the base of the anthers (see Fig. 2). Gradation is also observed in the locules that show an incomplete separation during development, sagittate locules and others with an irregular shape. Between four and eight glands were observed in L. ramiroi, some of them free and others fused at their bases, but always with clearly separated apices (Fig. 2). In L. ramiroi, the development of more than three fertile stamens in whorl III, with the corresponding increase in glands, could be a teratological condition due to an environmental response, possibly to the infection of Diptera larvae, which were observed parasitising some of the dissected flowers. To determine if the aberrations observed in the flowers of the species are due to the presence of the indicated parasites, further research will be required.
Licaria has been a well-defined genus by its three bisporangiate fertile stamens in whorl III, the presence or absence of staminodes in whorls I and II and the lack of staminodes in whorl IV, a deep receptacle and a double-margined cupule (Kurz 2000). Recent molecular studies have recovered a clade within the Ocotea complex called "Licaria group and allies", which includes, in addition to Licaria, other genera with deep receptacles and a double-margined cupule, including species of Ocotea Aubl., Aniba Aubl., Dicypellium Nees & Mart., Kubitzkia van der Werff, Paraia Rohwer, H.G. Richt. & van der Werff and Urbanodendron Mez (Trofimov et al. 2019).
The double-margined cupule in L. ramiroi suggests a morphological connection to species in genera such as Aiouea Aubl., Damburneya Raf. and Mespilodaphne Nees & Mart., especially with those with bisporangiate anthers. However, the species of these genera present other characteristics that are not recorded in L. ramiroi, such as the fact that, in Aiouea, there are staminodes in the fourth androecial whorl that show a large glandular head, generally chordate to sagittate (Rohde et al. 2017); in Damburneya, the tepals are adaxially pubescent at the base and generally papillose towards the tips (Trofimov et al. 2016) and both genera have nine fertile stamens, occasionally six or three in Aiouea. Mespilodaphne, on the other hand, is a recently reinstated genus, it presents flowers with spreading tepals, tongue-shaped stamens, heavily papillose and tetrasporangiate (Trofimov et al. 2019). Mocinnodaphne Lorea-Hern., recently included in Aiouea, meets most of the characters of Licaria, with the exception of having well-developed staminodes in whorl IV (Lorea-Hernández 1995). Due to a set of characters present in L. ramiroi, such as its deep receptacle, fertile stamens in the III whorl, bisporangiate, cupule with double margin, lack of staminodes in the first two whorls and the fourth, we consider that the best inclusion of the species is in Licaria.
Understanding the morphological complexity of the "Licaria group and allies" within the Ocotea complex and taking into account the androecial morphology recorded in L. ramiroi, considering that it is a genetic condition, then the species could be relocated to another genus within the Ocotea complex, but it would be necessary to wait for the species to be included in some molecular study to better define its phylogenetic relationships.